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A qualitative and quantitative assessment of seaweed decomposition in the Strait of Georgia Smith, Barry D. 1979-03-03

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A QUALITATIVE AND QUANTITATIVE ASSESSMENT OF SEAWEED DECOMPOSITION IN THE STRAIT OF GEORGIA by BARRY DOUGLAS SMITH B.Sc. (Honours), University of New Brunswick, 1974 A THESIS SUBMITTED IN PARTIAL FULFILMENT OF THE REQUIREMENTS FOR THE DEGREE OF MASTER OF SCIENCE in THE FACULTY OF GRADUATE STUDIES (Department of Botany) We accept this- thesis as conforming to the required standard THE UNIVERSITY OF BRITISH COLUMBIA June, 1979 (c) Barry Douglas Smith, 1979 In presenting this thesis in partial fulfilment of the requirements for an advanced degree at the University of British Columbia, I agree that the Library shall make it freely available for reference and study. I further agree that permission for extensive copying of this thesis for scholarly purposes may be granted by the Head of my Department or by his representatives. It is understood that copying or publication of this thesis for financial gain shall not be allowed without my written permission. Department The University of British Columbia 2075 Wesbrook Place Vancouver, Canada V6T 1W5 Date ABSTRACT Appropriate sampling and experimental programs resulted in a qualitative and quantitative assessment of seaweed litter biomasses, decom position rates and concomitant changes in nitrogen content; detritus biomass and decomposition rates; and faunal distribution patterns for the significant species within a successional seaweed community in the Strait of Georgia, British Columbia, Canada. A simulation model incorporating suitable data obtained from these sampling and experimental programs facilitated prediction of detritus formation rates, biomass, nitrogen content and the seasonal availability of detritus as a food resource for fauna. Soluble matter release rates from decomposing seaweed litter and its nitrogen content were also determined. Of the ca 43 taxa identified within the seaweed litter collec tions, Fucus distichvs L. (41%), Irldaea cordata (Turner) Bory (26%), Nereocystis 1 lEtkeana (Mertens) Postels and Ruprecht (27%), and Laminaria (4%) (L. saccharina (L.) Lamouroux and L. groenlandica Rosenvinge) accounted for more than 97% of total litter deposition. The mean peak summer biomass of all litter was ca 5 g 2 ash-free dry weight (AFDW)/m with this figure approaching zero during January and February. Litter distribution was patchy and there was sufficient evidence to conclude that most litter was retained, and underwent decomposition, in the immediate vicinity of its place of deposition. Litter decomposition experiments performed on the 10 most signi ficant contributors to seaweed community structure indicated that decomposition of seaweed litter occurs rapidly compared to vascular plant litter. The time required for seaweed litter to disappear from 2 rrrn mesh litter bags ranged from six days, for the lamina of Nereocystis lnetkeana, to ca 70 days, for Fucus distichus . Some similarity in decomposition rates was observed amongst species displaying taxonomic and/or morphologic affinities. Assessment of nitrogen content of decomposing seaweed litter revealed that nine of the 10 species assayed lost nitrogen less rapidly than total litter biomass. As determined by assaying microbial consumption of particulate material, the time required for detritus (particle size < 1 mm, dry) to fully decompose was short. Of the 10 species tested, Iridaea cordata detritus decom posed most rapidly at a rate of 5.7% per day while rates for Gigartina papillata (C. Agardh) J. Agardh, Laminaria groenlandica, Laminaria saccharina and Nereocys-tis luetkeana ranged from 2-4% per day. Data for the remaining species were less conclusive although all decomposed at rates less than one percent per day. Variation in specific decomposition rates was shown to be correlated with the structural composition of the detritus. Those species with a relatively small percentage of crude fibre as a component of their particulate fraction decomposed more rapidly than those species with a higher percentage of crude fibre. For the two most rapidly decomposing species, Iridaea cordata and Nereocystis luet keana, a trend toward a more rapid decomposition rate as mean particle size decreased was evident. Natural detritus (particle size < 2 mm, wet) biomass accumulation 2 within the study site peaked at ca 1.4 g AFDW/m during the latter half of August 19 76. This value represents 1-5% of the quantity of detritus predicted to have been formed from seaweed litter alone and a lesser percentage of the total quan tity of seaweed detritus formed. Exportation out of the seaweed zone is believed to be responsible for this discrepancy. The predicted rates of detritus forma tion and soluble matter release from decomposing seaweed litter peaked at ca 0.6 2 and 0.5 g AFDW/m per day, respectively, in early September 1976 from a low near zero in February. In total, ca 56% of litter biomass formed detritus, the re mainder being released as soluble matter. The mean nitrogen contents of the detritus formed and the soluble matter released were 2.48 ± 0.03% and 1.36 ± 0.03 of their dry weights, respectively. The annual contribution of seaweed litter biomass via detritus and soluble matter to local coastal waters is estimated to 2 be in the range of 70-85 g C/m . Detritus formed from seaweed litter was determined to have a C:N ratio of 10-13:1, rendering it suitably nutritious for utilization by fauna as a food resource, however it could not be shown conclusively that the coincidence, en masse, of specific fauna and maximum detritus availability was a response to the availability of detritus as a food resource. The possibility of such a correlation is discussed with reference to two species of caprellids, Caprella alaskana Mayer and Metacaprella anomala Mayer, and the benthic gastropod Lacuna marmorata Dall. iv TABLE OF CONTENTS ABSTRACT LIST OF TABLES vii LIST OF FIGURES ix ACKNOWLEDGEMENTS '. xi INTRODUCTION 1 METHODS The Study Area 7 Sampling Litter assessmentDetritus assessment 10 Faunal assessment 1 Field Experiments Litter decomposition experiments 12 Litter senescence experiments 3 Laboratory Experiments Detritus decomposition Experiment 1 (microbial oxygen consumption) 15 Experiment 2 (microbial consumption of particulate material) 15 Nitrogen content of decomposing litter 17 Structural composition of species contributing to litter .... 17 Model Development and Data Analysis 19 RESULTS Litter assessment 20 Structural composition of species contributing to litter .... 40 Litter decomposition experiments 42 Litter senescence experiments 54 Nitrogen content of decomposing litter 5Detritus decomposition Experiment 1 (microbial oxygen consumption) 57 Experiment 2 (microbial consumption of particulate material) 62 Detritus assessment 7Faunal assessmentDISCUSSION Litter assessment 80 Litter decomposition experiments 82 Nitrogen content of decomposing litter 85 Detritus decomposition • 86 Detritus assessment 91 Faunal assessment 2 v SIMULATION MODEL OF LITTER AND DETRITUS PROCESSING Introduction 98 Model development 99 Results 105 Discussion Ill SUMMATION 117 LITERATURE CITED 119 APPENDICES I Litter assessment data 12 7 II Faunal assessment data 144 III Detritus assessment data 153 IV Depth data 156 V Litter decomposition experimental data 157 VT Detritus decomposition data (Experiment 1) 159 VTI Detritus decomposition data (Experiment 2) 160 VIII Simulation model computer program 161 vi LIST OF TABLES Table 1: Mean biomass per m of the major contributors to the litter pool within Site 1 based on the collections of 27 July and 3 August 1976. 21 Table 2: Comparison between the number of living Nereocystis luet-keana plants within a transect belt and the quantity of Nereocystis luetkeana litter collected within the same belt. 28 Table 3: Comparison of the total quantity and specific composition of litter collected within the transect at 95 m within Site 1 on 9 November 1975 and the total quantity and specific com position of litter collected within the transect at Site 2 on 10 November 1975 (gr AFDW/transect) . 31 Table 4: The percentages of each of the soluble, moderately resistant and crude fibre components of the significant species within Site 1. 41 Table 5: Number of days required for living portions of the major contributors to the litter pool within Site 1 to leave a 1.0 cm mesh litter bag under shaded and exposed conditions, 55 Table 6: Percentage nitrogen content of the material remaining with in the litter bags at the termination of their incubation period. 56 Table 7: Analysis of variance table for the results of Experiment 1, demonstrating the effects of particle size, detrital species and length of incubation period on the oxygen consumption by microbes utilizing the detritus as a carbon source. 59 Table 8: a) Subsets delimited by Duncan's New Multiple Range Test. Each subset contains those detrital species which show a signifi cant (p < .05) degree of affinity with respect to the quanti ty of oxygen consumed by microbes decomposing the detritus, b) The average percentage soluble content of the subsets in Table 8a. 63 Table 9: Analysis of variance table for the results of Experiment 2, demonstrating the effects of particle size, detrital species and length of incubation period on the consumption of parti culate material by microbes utilizing detritus as a carbon source. 65 Table 10: Subsets delimited by Newman - Keul's Range Test. Each sub set contains those detrital species which show a significant (p < .05) degree of affinity with respect to the quantity of particulate material consumed by microbes decomposing the detritus. 68 vii Table lib: Table 11a: The total number of each faunal species summed over the 28 July, 18 August and 12 September 1976 transect collec tions. The percentage that this number represents of the total number of occurrences over the entire sampling period is in parentheses. 74 The total dry weight of each faunal species summed over the 28 July, 18 August and 12 September 1976 transect collections. The percentage that this figure represents of the total dry weight of individuals collected over the entire sampling period is in parentheses. 75 Table 12: History of the occurrence (per m?) of two species of Cap-rellidae, Caprella alaskana and Metacaprella anomala, with in the summer faunal collections of Dr. R. E. Foreman (un published) . 95 Table 13: Mean monthly temperatures (a) and the corresponding decom position rate adjustment factor (b) for the period November 19 75 until October 1976. 102 Table 14: Comparison of the percentage contributions by the major contributors to the litter pool within Site 1 as deter mined by litter biomass alone and application of the de composition rates of these species to litter biomass data. 10 7 viii LIST OF FIGURES Figure 1: Figure 2: Figure 3: Figure 4: Figure 5: Figure 6: Figure 7: Figure 8: Figure 9: Location of field study sites. 8 Spatial characteristics of litter biomass for the major contributors to the litter pool within Site 1 based on the collections of 27 July and 3 August 1976. 22 Distribution of Laminaria litter collected along the tran sect at Site 2 on 10 November 1975 relative to depth below mean sea level. 9 Depth contours (zn below mean sea level) for Site 1. 32 Seasonal distribution of litter biomass for the major con tributors to the litter pool within Site 1 based on collec tions along the 95 m transect location at 3-4 week intervals for the period 20 August 1975 until 2 October 1976. 33 Litter decomposition curves (submodels) calculated from data obtained in the litter bag experiments. 4 3 Plot demonstrating an increase in the ratio of nitrogen: dry weight biomass of decomposing litter relative to unde-composed litter. 58 Cumulative oxygen consumption by microbes decomposing the 10 detrital species in Experiment 1. 61 Relationship between the percentage soluble contents of the 10 detrital species (exclusive of Iridaea cordata) and the quantity of oxygen consumed by microbes decomposing the detri tus after five days of incubation, as determined in Experiment 1. 64 Figure 10: Cumulative loss of particulate material from the 10 detri tal species decomposed in Experiment 2. 67 Figure 11: Cumulative loss of particulate material from Iridaea cor data and Nereocystis luetkeana (stipe and lamina combined) detritus. For each species the results for the three detri tal particle sizes are presented. 70 Figure 12: Relationship between the maximum percentage loss of particu late material from the 10 detrital species decomposed in Experiment 2 and the percentage of crude fibre in the parti culate material of each detrital species. 71 Figure 13: Contour representation of detritus biomass along the 95 m transect location within Site 1 for the period 28 May until 7 October 1976. 73 Figure 14: Seasonal distribution histograms of the total number and dry weight (g) of Cancer oregonensis, Metacaprella anomala and Lacuna marmorata occurring within the seven transect collec tions from 25 May until 7 October 1976. ix Figure 15: Seasonal trend in the mean dry weight (g) per individual of Lacuna marmorata for the period 25 May until 7 October 19 76. 78 Figure 16: Spatial distribution along the 95 m transect location with in Site 1 of Lacuna marmorata (numbers and biomass) and detri tus biomass demonstrating a coincidence in the occurrence of their maximum abundances. 79 Figure 17: Tenth degree polynomic curve fitted to the seasonal biomass data obtained from litter collections along the 95 m tran sect location within Site 1 from 20 August 1975 until 2 October 1976. 101 Figure 18: Flow chart outlining the major operations involved in the simulation of litter and detritus processing within Site 1. 106 Figure 19: Seasonal profiles for the formation rate of detritus and the release rate of soluble matter from decomposing sea weed litter biomass within Site 1. 108 Figure 20: Detritus biomass predicted for the 95 m transect location' within Site 1 based on litter collections from that loca tion only. 109 Figure 21: Detritus biomass predicted for Site 1 based on litter col lections from all transect locations within Site 1. 110 x ACKNOWLEDGEMENTS Many persons need to be credited for their contribution to ward the successful completion of this thesis. I appreciate the supervision of Dr. Ronald E. Foreman who provided research facilities and first hand ex perience with the study of marine macrophyte systems. Mr. Thomas Nicol was very helpful with computing problems, particularly during development of the simulation model. Perhaps the most important contribution to this work was the effort of my SCUBA partners. I am especially grateful to Mr. Eric L. Cabot who was my diving buddy for most of the field sampling exercises. The staffs of the Woodward Biomedical Library and the U.B.C. Computing Centre provided excellent service. I thank Ms. Nancy A. Smith for her as sistance with the typing. xi - .1 -INTRODUCTION Primary production by terrestrial and aquatic plants is the major source of food energy for consumer organisms. In many cases it has been shown that heterotrophic utilization of primary production involves largely a delayed consumption of detritus. Darnell- (1976a) defines detritus as being "all types of biogenic material in various stages of microbial decom position which represent potential energy sources for consumer species". This definition is appropriate, but includes material that this study interprets as 'litter', defined as larger, less fractured material whose biogenic origin can be easily recognized. The importance of detritus as a food source for consumers has been demonstrated for several ecosystem types. In an east coast salt marsh studied by Teal (1962) only 7% of the net primary production was utilized in herbivore respiration while 47% was utilized by decomposer organisms associated with detritus derived from Spartina litter. Similarly, data for several terrestrial systems indicate that 62-100% of net primary production enters the litter pool (Rodin and Bazilevich 1967) with future processing forming detritus. An exception to this trend is found in plankton based systems where Up to 90% of the primary production may be consumed by zooplankton grazers. In such cases a large portion of the material consumed may pass through the gut of the zooplankters without being assimilated, and enter the decomposer food chain. This is especially true during bloom conditions (Cushing 1964). To date, studies concerning detritus formation and utilization in coastal marine ecosystems have dealt mainly with aquatic vascular plants such as Zostera marina L. (Harrison and Mann 1975 a&b, Harrison 1977, Tenore et al.1977), Thalassia testudinum Banks ex Konig (Fenchel 1970, Wolff 1976, Knauer and Ayers 1977) , mangroves (Heald 1969) and Spartina alterniflora Loisel as well as other salt marsh plants (Odum and de la Cruz 1967, de la Cruz - 2 -and Gabriel 1974, Gosselink and Kirby 1974, de la Cruz 19 75, Gallagher et al. 1976, Pickral and Odum 1976, Hanson and Weibe 1977). This work has been re viewed by Fenchel (1972, 1973). The importance of associated microorganisms in this process has been stressed by Johannes' (1965), Seki (1972), Fenchel and Harrison (1976), and Heinle et al. (1977). These studies have been largely of a qualitative nature with little attempt to quantify plant detrital contribu tions to coastal energy flow. There are but a few studies concerning detritus formation by attached marine macrophytes. Although estimates of primary production for the coastal seaweed zone indicate that these areas are amongst the most highly pro ductive in the world (Clendenning 19 71, Mann 19 72a) very little is known of the fate of this production. With the macrophytic fringe of the oceans having a productivity that may be up to 40 times that of the ocean (Mann 19 72a) and a standing crop exceeding that of phytoplankton by 100 fold (Blinks 1955) , the possibility of its having a more than token contribution to the energy flow in near-shore ecosystems of which some commercial fish species may be components becomes a reality. Salmon (Oncorhynchus spp.) and herring (Clupea harengus pallasii Valenciennes), currently the most valuble fish to the British Columbia economy (Statistics Canada 1976) spend critical times of their lives in near-shore waters. Herring are dependent on seaweed as substrate for their spawn (Taylor 1964). Young salmon feed in estuarine waters (Sibert et al.1977). Mann (1972b) estimates the yearly productivity of the seaweed 2 zone in St. Margaret's Bay at 1750 g C/m . This makes the seaweed zone the only primary marine resource with a confirmed yearly production greater than 1 kg C/m2. Possible fates of seaweed production are: 1. exudation as soluble matter - 3 -2. consumption by herbivores 3. erosion and fragmentation from lamina tips 4. release as reproductive structures 5. natural mortality 1) The release of soluble organic compounds from marine sea weeds was first demonstrated by Craigie and MacLachlan (1964). Later Sieburth and Jensen (1968) and Sieburth (1969) established that exudation from marine macrophytes is comparable to that of phytoplankton which Fogg (1966) states to lie between 5% and 35% of total carbon fixed within a population. Fucus vesi-culosis L. was estimated to lose 30.7% of its total carbon budget as exudate, at an average rate of 41.6 mg C/100 g/hr. These rates are comparable to those obtained for other Phaeophyta; 44.6, 37.8 and 31.3 for Laminaria digitata (L.) Lamouroux, Laminaria agardhii Kjellman, and Ascophyllum nodosum (L.) Le Jolis, respectively. Chondrus crispus Stackhouse (Rhodophyta) was significantly lower at 4.4 mg C/100 g/hr. Johnston et al. (1977) determined that up to 36% of total carbon fixed by Laminaria saccharina (L.) Lamouroux was released extracellularly. Brylinsky (1977) examined two species each of Rhodophyta, Acanthophora spicifera (Vahl) Borgesen and Chondria dasyphylla (Woodward) C. Agardh, and non-kelp Phaeo phyta, Dictyota dichotoma (Hudson) Lamouroux and Sargassum natans (L.) Meyen, and determined physiological release rates of less than 4.0% of total carbon, disclosing an apparent disparity in release rates between kelp-like seaweeds and others. 2) Sea urchins are generally recognized as the most significant and prominent grazers in temperate seaweed systems. Miller and Mann (1973) concluded that the green sea urchin Strongylocentrotus droebachiensis Muller, the apparent major herbivore in eastern Canada, consumed only 1-7% of seaweed net production during their period of study. With Strongylocentrotus droebachi ensis accounting for 80% of the herbivory in'this area (Miller et al. 1971), - 4 -consumption of seaweed biomass approached 10% of net production. On occasion large numbers of sea urchins have severely per-turbated the seaweed zone (Leighton et al. 1966, Paine and Vadas 1969, Leighton 1971, Miller and Mann 1973, Breen and Mann 1976, Foreman 1977, Mann 1977). This has sometimes resulted in total denudation of the affected area both by direct grazing and by detaching plants from the substrate. Dur ing these periods the detached plants complement dead plant material from other sources in contributing to the pool of marine plant litter. 3) Johnston et al. (1977) present quantitative information on erosion from lamina tips. They estimate that for Laminaria saccharina growing in a sheltered location near the head of Loch Creran, Scotland, 40-50% of annual gross production is lost by distal decay, resulting in a contribu tion to either the detrital or litter pools depending on whether the loss is via erosion or fragmentation, respectively. Plants growing in more exposed locations might be expected to lose a higher percentage of their carbon budget by distal decay. Laycock (1974) demonstrated that large populations of bacteria associated with the lamina tips of Laminaria longicruris de la Pylaie were at least partially responsible for distal decay. 4) As release of reproductive structures would be indistin guishable from the exudation of soluble matter or loss of particulate biomass, the need to consider reproductive losses separately is precluded. 5) Natural mortality constitutes the final exit pathway. The death of the seaweeds initiates their entry into the pool of marine plant litter where they undergo decomposition concomitant with the formation of detritus and detritus processing. In an attempt to place the various aspects of the seaweed 'biomass budget' into perspective, Khailov and Burlakova (1969) proposed a - 5 -quantitative partitioning of the total gross production of seaweeds into suitable compartments. From experiments with five species of Barents Sea mac-rophytes and 13 species of Black Sea macrophytes they judge loss due to con sumption by herbivores to be ca 11.2% and calculate that 37.3% of gross pro duction is represented by living biomass, the major source of detritus, either via erosive or litter pathways. With the realization that the contribution of seaweed pro duction to the detrital pool may exceed its consumption by herbivores by three to four fold it does not seem unreasonable or premature to suggest that detritus processing is an essential aspect of energy flow in near shore ecosystems. To confirm this hypothesis it is necessary that the dynamics of seaweed litter decomposition along with subsequent detritus formation, processing, and utiliza tion be investigated. This thesis descriptively and quantitatively assesses the con tribution of seaweed litter biomass to the detrital pool. The objectives of the study were: 1) to determine the total quantity and seasonal abundance of seaweed litter available as a source of detritus in a defined area 2) to determine the formation rate, longevity and decomposition rate of detritus formed from selected seaweed species 3) to predict the seasonal rates of detritus formation, its biomass and nitrogen content for a defined area, and assess its impor tance as a food resource for fauna 4) to characterize selected seaweed species in terms of their 'soluble', 'moderately resis tant', and 'crude fibre' components and cor relate differences in the relative quantities of these components with observed decomposi tion rates for litter and detritus. - 6 -These objectives were realized by conducting specific samp ling and experimental programs and by execution of a simulation model of litter and detritus processing based on data acquired from these programs. - t -METHODS THE STUDY AREA All field work was carried out in the shallow sublittoral zone adjacent to the southeastern shore of Bath Island, British Columbia, the east ern most of a cluster of small islands known as The Flat Tops. These islands are ca 32 km west of the mouth of the Fraser River, and hug the southeastern extension of Gabriola Island, the northernmost of a group of islands called The Gulf Islands (Figure 1). Bath Island is 3.2 hectares in area, its main geological component being sandstone complemented with minor amounts of shale and conglomerate (Muller 1971). The main research area is a gently sloping one hectare plot well exposed to the southeast. The plot can be appropriately des cribed as a successional kelp bed due particularly to the extensive stand of Nereocystis luetkeana (Mertens) Postels and Ruprecht which does well there (Foreman .1977). This one hectare plot will be known as Site 1. A second loca tion, near Site 1, will be referenced as Site 2. All laboratory work was performed in the Department of Botany at the University of British Columbia. SAMPLING Three sampling programs were implemented: 1) to determine the seasonal and spatial distribution of seaweed litter biomass within Site 1 2) to determine the seasonal distribution of detritus biomass within Site 1 3) to determine the seasonal and spatial distribution of invertebrate fauna within Site 1. Litter Assessment: The main, permanently marked transect location intersected the shore at 95 m along the 100 m shore front forming the base of Site 1. At times - 8 -Figure 1. Locations of field study sites. a) Fiat Top Islands in relation to the lower mainland of British Columbia. b) Site 1 and Site 2 in relation to the Flat Top Islands. of sampling a line 100 m in length was extended from the high intertidal zone (upper limit of barnacles) to a point beyond the zone of most seaweed cover. No significant accumulations of litter were observed outside of the zone sampled. Two scuba divers then proceeded to collect all seaweed litter that lay within a metre on either side of the transect line. The transect was segmented into ten 20 m2 quadrats with the collections from each being placed in an appropriately labelled bag. On occasion, when the quantity of litter within a standard 20 m2 quadrat was more than could be easily collected, the quadrats were subsampled in a representative fashion. Sampling at this site was carried out at ca 3-4 week intervals from August 1975 until October 1976. These data were used to determine the seasonality of the biomass of seaweed lit ter. On 3 August 1976 similar transects were sampled from 5, 35 and 65: m along the base in order to determine the spatial distribution of litter within Site 1. On one occasion (10 November 1975) a single transect was collected at Site 2, ca 200 m away and less exposed than Site 1, allowing a comparison of the two areas to be made. When collections were made a seaweed was classified as litter if it could be described by one of the following phrases: 1) detached and having settled to the bottom, generally snagged amongst rocks or debris 2) attached but apparently dead 3) in the case of Nereocystis luetkeana stipes, attached or unattached and lying prone, the pneumatocyst having flooded. For each site a transect depth profile was recorded and for each quadrat the substrate was described. On 3 August 1976 the number of living Nereocystis luetkeana plants in each quadrat of the four transects located - 10 -within Site 1 was enumerated. All collections were transported to the laboratory where they were sorted and identified as precisely as possible according to Widdowson (1973, 1974) and Lindstrom et al. (1974). Laminaria saccharina and Laminaria groenlandica were not always distinguishable and so were often recorded only as Laminaria. Nereocystis luetkeana was subscripted as either stipe or lamina litter. For each taxon in every quadrat the wet weight, dry weight (24 hours at 100 C) and ash-free dry weight (12 hours at 425 C) were recorded. Detritus Assessment: From May 1976 until October 19 76 at ca three week intervals the biomass of detritus within Site 1 was determined. Nine permanent qua drat locations were fixed, roughly corresponding to 20,30... 100 m along a transect perpendicular to the shore at 95 m along the base of Site 1. The actual positioning of the quadrat was determined by the availability of rela-2 tively flat, continuous substrate extensive enough to accommodate a 0.0625 m quadrat. Each of these quadrat locations was initially scrubbed clean with a wire brush, and again following each sampling period. Detritus was collected using a hand pump designed for bailing small boats. It was modified by securing an 11 lb plastic bag to the exhaust port. By operating the pump in a normal fashion, passing the intake port over the quadrat, all loose material was sucked into the bag. Control samples were collected by drawing sea water into the bag while the intake port was well above the substrate. Upon returning to shore, the contents of each bag were screened through 2 mm mesh household screening to remove large particles, then passed through preweighed Whatman GF/C^glass fibre filters (2-3 pm pore size) using a Millipore® filter apparatus. The residuum was dry weighed (12 hours at 100 C) and ash-free dry weighed (4 hours at 42 5 C). - 11 -Faunal Assessment: From May until October 1976 at approximately three week intervals, faunal collections were made within Site 1. The sampling procedure involved the 2 collection of 0.0625 m quadrats at 30,40...100 m along the permanently located transect at 95 m along the base of Site 1. The organisms were collected using an underwater airlift (Foreman 1977) and trapped in a collecting bag made from panty hose. Samples were transported to the laboratory while fresh where they were sorted, identified according to Kozloff (1974), counted, and wet and dry (24 hours at 100 C) weighed. FIELD EXPERIMENTS Two field experiments were performed during July and August 19 76. The first was designed to obtain in situ rates of decomposition for killed sea weeds, the second to estimate senescence times for those species contributing significantly to the litter within Site 1. Litter Decomposition Experiments: Seaweeds were chosen for the litter decomposition experiments on the basis of their contributions to the standing crop of living seaweed biomass within Site 1 (coralline algae excluded). As Site 1 overlaps almost entirely the one hectare plot Foreman (1977) defined for his biomass studies in 1972, his data were used as a criterion for ranking the seaweeds. They are, in de-cending order of their 'importance values' (Foreman unpub.) : Iridaea cordata (Turner) Bory Constantinea subulifera Setchell Laminaria (L. saccharina, L. groenlandica Rosenvinge) Fucus distichus L. Odonthalia floccosa (Esper) Falkenberg Rhodomela larix (Turner) C. Agardh Plocamium coccineum var. pacificum (Kylin) Dawson Gigartina papillata (C. Agardh) J. Agardh Nereocystis luetkeana The above species accounted for just over 80% of seaweed standing crop biomass, exclusive of coralline algae, as their contribution to litter would be minor. In the litter bag experiments Laminaria saccharina and Laminaria groenlandica were considered separately as were the stipe and lamina sections of Nereocystis luetkeana, bringing the total count of individual experiments to 11. The appropriate seaweeds were collected live, cut into portions suitable for the litter bags, wet weighed and killed by placing them in a seawater bath at ca 50 C for 10-15 minutes. A separate portion of each seaweed, a control, was wet and dry weighed without undergoing decomposition. - 13 -The remaining portions were placed in 15 cm x 15 cm litter bags made from plastic household screening (2 mm mesh). Three litter bags were pre pared for each seaweed tested with the exception of Fucus distichus for which four bags were prepared. Each litter bag was placed in a larger (1 cm mesh) bag and suspended from the mesh (5 cm) forming the roof of an aluminum framed cage (2.0 m x 1.5 m x 0.5 m) constructed as a precaution to reduce the inter ference of large animals which might graze upon or otherwise interact with the decomposing seaweed. The cage was placed on the bottom at ca 6 m below mean sea level in a relatively sheltered embayment (Site 2). From preliminary experiments it was judged that the breakdown of the seaweeds would be rapid, therefore the litter bags were retrieved based on visual observations of the progression of the decomposition process rather than according to a predeter mined schedule. The material which remained in the litter bags at the termina tion of the incubation period was removed, dry weighed and saved for nitrogen determination. Following completion of all incubations the dry weights were normalized with respect to the control and expressed as a percentage of the original dry weight of the material placed in the litter bags. Litter Senescence Experiments: A second experiment was performed to determine the time required for the seaweeds which appeared to be the more significant contributors to the litter within Site 1 to die once having entered the litter pool. Death is con sidered to be the time when tissue breakdown by autolytic or saprophytic means begins. The species chosen were Nereocystis luetkeana (stipe and lamina sec tions) , Laminaria saccharina, Laminaria groenlandica and Iridaea cordata. Live portions of each of these seaweeds were placed in 1 cm mesh litter bags (not necessarily a single species per bag) and secured to the substrate within Site 1 at ca 3-5 m depth. Some bags were left exposed while others were placed between rocks or within shaded crevices. These bags were observed over five weeks, - 14 -noting changes in the condition of their contents. The time required for a seaweed to die was estimated by as suming that once dead, the number of days required for seaweed biomass to leave a 1 cm mesh litter bag was about one half the number of days required for it to leave a 2 mm mesh bag. The latter data are known from the litter decomposition experiments. By subtracting the latter number of days from the number of days required for the unkilled seaweed to disappear from the 1 cm mesh bags, the length of time required for fresh litter to die was esti mated. - 15 -LABORATORY EXPERIMENTS Detritus Decomposition: Detritus was created from seaweed species which had been collec-® ted live, washed, cleaned, dried, crushed by hand and processed in a Wiley mill. Three size fractions of detritus (1000-420 ym, 250-149 ym and 44-0 pm) q were then collected by shaking the crushed seaweed through a series of Endicott sieves. The ratio of surface area exposed to microbial attack for the three size categories will be, from the largest to the smallest, ca 1:4:32, when all are present in equal mass. By setting the upper limit of detrital particle size at 1.0 mm (dry) the detritus decomposition experiments can be considered a continuation of the litter decomposition experiments which assessed the forma tion rate of detrital particles < 2.0 mm (wet). The detritus was derived from the same 10 species used in the litter bag experiments, the stipe and lamina sections of Nereocystis luetkeana being considered separately. Two experiments were performed to assess the microbial utiliza tion of this detritus, one based on oxygen consumption, the second based on microbial consumption of particulate material. Both experiments were structured around a 3 x 3 x 11 factorial design (Hicks 19 73) incorporating three particle sizes, three incubation periods, and 11 experimental sets (10 species). Experiment 1 (Microbial Oxygen Consumption): Assessment of oxygen consumption required 12 acid-washed, 300 mL Biochemical Oxygen Demand (BOD) bottles for each incubation set. Oxygen content was assayed by the Winkler method (Strickland and Parsons 1972) . Into three of each subset of four bottles, a 1.0 mg plug of detritus of a single size class was placed; the fourth remained a control. This procedure was repeated for the other two size classes. An inoculum of 1.0 mL of fresh seawater was pipetted into each BOD bottle as a source of microbes, following which all bottles were -in filled with filtered (0.45 ym) and aerated seawater. The bottles were capped and incubated in a 15 C water bath and agitated daily. Bottles represen ting each particle size, and a control (four in total) were removed after each of 5, 10, and 20 days of incubation. They were immediately fixed with the appropriate reagents. Twenty days was sufficient time to allow a significant drop in the oxygen content of the bottles while avoiding depletion. This procedure was repeated for all 11 sets. Experiment 2 (Microbial Consumption of Particulate Material) .-Each incubation set for the experiments to assess loss of parti culate matter required eighteen 250 mL Erlenmyer flasks (six flasks per size class) as culture vessels. A 0.1 g plug of detritus representing a single size class was added to each flask. These 18 flasks were divided into two equal sets. To one set, the control, 100 mL of 0.45 pm filtered, sterile seawater containing KCN at a concentration of 0.1% (Harrison and Mann 1975b) was added. The second set received 100 mL of the sterile seawater enhanced with 0.15 g/L of NaNO^ (Gosselink and Kirby 1974) and was inoculated with 1.0 mL of fresh seawater. Each experimental flask was thus paired with a control flask. All flasks were incubated at 15 C and agitated regularly. That sterility prevailed in the con trol flasks was confirmed by the clarity of the control flasks when compared to the experimental flasks. At 10, 20, and 30 day intervals an experimental flask and a con trol flask of each particle size (six in total) were retrieved. The contents of ® each flask were filtered through preweighed Whatman GF/C glass fibre filters. The filters were dried (4 hours at 100 C) and weighed. The loss of particulate ma terial for any treatment group was determined by subtracting the residue weight for each experimental flask from that of the control flask. Nitrogen Content of Decomposing Litter: The total nitrogen content of the seaweed material which re mained in the litter bags at the time they were retrieved was determined using a macro-Kjeldahl method (Skoog and West 1969). The quantity of nitrogen obtained in each assay was expressed as a percentage of the total dry weight of the mater ial assayed. Structural Composition of Species Contributing to Litter: For all 10 seaweed species and the stipe and lamina sections of Nereocystis luetkeana the contribution by each of three basic structural com ponents to living seaweed biomass was determined on a dry weight basis. These components will be referred to as the 'soluble', 'moderately resistant' and 'crude fibre' components. For experimental purposes material which passed through a filter of 2-3 pm pore size was classified as 'soluble'- 'Moderately resistant' refers to material which is particulate and easily metabolized by microbes, being com posed largely of low molecular weight and non-structural polymeric compounds within the cell matrix. 'Crude fibre' consists mainly of cellulosic sugar poly mers that are somewhat resistant to the attack of microbes. These polymers are generally responsible for the structural integrity of cell walls (Steward 1974). Both the soluble and crude fibre components were determined ex plicitly. The quantity of soluble matter was determined in Experiment 2 of the detritus decomposition experiments. The weight of the residuum obtained from filtering (2-3 ym pore size) the contents of the control flasks at the end of each incubation period was subtracted from the initial weight (0.1 g) of the material in the flasks, this being the quantity of material passed through the filter, i.e. the soluble content. Accepted values for soluble content were ob tained by averaging the results of the 10 and 20 day incubation periods since - 18 -by day 30 there were indications that some of the control flasks were no longer sterile. An analysis was performed using the method described by Strickland and Parsons (1972) to determine the percentage of crude fibre present in seaweed biomass. The dry weight of the crude fibre fraction was determined following extraction of the alkali/acid-soluble components of 30 mg samples of ground seaweed (0-44 urn particle size). Crude fibre carbohydrate content (expressed as an equivalent amount of glucose) was determined spectrophotometrically. Sample sizes were 1.0 mg. - 19 -MODEL DEVELOPMENT AND DATA ANALYSIS Much of the data required from the previously described sam pling and experimental programs were suitable for incorporation into a mathe matical model created to simulate the transport of decomposing seaweed biomass through detrital pathways. Most of the data were acquired with this end in mind. The model also incorporated environmental data measured during 1975 and 1976 as a part of an ongoing program by Foreman (unpublished) to describe the meteorological and oceanographic conditions of the area. The model was written in FORTRAN G and debugged and executed by the IBM 370 computer at the University of British Columbia Computing Centre. In addition, numerous support programs and subroutines were used in the analysis of experiments and presentation of results. -3.0-RESULTS Litter Assessment: Five species (four genera) of seaweeds were responsible for more than 97% of the plant litter collected over the 14 month sampling period from 20 August 1975 until 2 October 1976. These species were Fucus distichus, Iridaea cordata, Nereocystis luetkeana, Laminaria saccharina and Laminaria groenlandica. In all, about 43 taxa were recognized within the litter collec tions. Table 1 summarizes the distribution of litter biomass collected from the transects at 5, 35 and 65 m within Site 1 on 3 August and at 95 m on 2 7 July 1976. These transects will be referred to collectively as the midsummer litter collections. Figure 2 (a-e) presents spatial representations of the distribu tion of litter at midsummer of 1976, near the time of maximum litter accumula tion. The area defined by the abcissa and ordinate represents Site 1 as though it were being observed from above. Note that the litter derived from Fucus distichus and Iridaea cordata, whose normal habitats are the intertidal and upper subtidal zones, respectively (Lindstrom 1973), is retained almost exclu sively within the shallow subtidal zone. Nereocystis luetkeana and Laminaria litter is retained in deeper water, in the zone where these plants grow ^abun-dantly. Table 2 demonstrates a positive correlation between the number of living Nereocystis luetkeana plants observed during each of the midsummer col lections and the quantity of Nereocystis luetkeana litter within these same collections, indicating that litter tends to be retained where it was deposited. From Figure 3 it can be seen that Laminaria litter at Site 2 was collected almost entirely within the outer extent of the transect, in a depth range of 4-5 m below MSL. This range is comparible to the kelp community zone delimited by Lindstrom (1973). Visual examination of the area confirmed a large standing crop biomass of Laminaria in the vicinity of Site 2 and within this depth range. Table 1. Mean biomass per m of the major contributors to the litter pool within Site 1 based on the collections of 27 July and 3 August 1976 • Species Wet weight (%1 Dry weight (%1 Ash-free dry weig Fucus distichus 27.3 (65.8) 5.40 (70.3) 3.96 (72.0) Iridaea cordata 4.6 (11.1) 1.20 (15.6) 0.83 (15.0) Nereocystis luetkeana (stipe) 1.2 ( 2.9) 0.16 ( 2.1) 0.11 ( 2.0) Nereocystis luetkeana (lamina) 6.3 (15.2) 0.62 ( 8.1) 0.41 ( 7.5) Laminaria 0.88 (2.1) 0.13 ( 1.7) 0.09 ( 1.6) All other species 1.18 ( 2.8) 0.17 ( 2.2) 0.11 ( 2.0) TOTAL 41.46 7.68 5.51 - 22 -Figure 2. Spatial characteristics of litter biomass for the major contributors to the litter pool within Site 1 based on the collections of 27 July and 3 August 1976. Contour intervals are in g ash-free dry weight per 10 IT?. Solid circles indicate pockets of litter. Contour interval a) Fucus distichus as labelled b) Iridaea cordata s labelled c) Nereocystis luetkeana (stipe) 1.0 d) Nereocystis luetkeana (lamina) 4.0 e) Laminaria as labelled - 23 -a) Fucus distichus o a 143 68 M 56 M 1261 1 1 1 1 1 0.0 20.0 40.0 60.0 80.0 100.0 DISTRNCE ALONG SHORE M) - 24 -b) Iridaea cordata a LU (Da CCS"1 cr: l— CD —I • , LU CJ -z. cr i— co°. •—io_| 16 51 227 1 1 1 I 1 0.0 20.0 40.0 60.0 80.0 100.0 DISTANCE ALONG SHORE (M) - -c) Nereocystis luetkeana (stipe) - 26 -e) Laminaria - 28 -Table 2. Comparison between the number of living Nereocystis luetkeana plants within a transect belt and the quantity of Nereocystis luetkeana litter collected within the same belt. Transects at 5, 35, 65 and 95 m along the base of Site 1 were collected either on 27 July or 3 August 1976. The transect at Site 2 was collected on 10 November 1975. Number of living Nereocystis luetkeana (per transect)  Site 1: 05 14 35 7 65 9 95 38 Site 2: 0 Quantity of Nereocystis luetkeana litter collected (g AFDW/transect) Stipes Lamina Total 4.21 32.04 36.25 6.18 27.47 33.65 4.55 26.17 30.72 27.90 76.57 104.47 2.71 4.09 6.80 Figure 3. Distribution of Laminaria litter collected along the transect at Site 2 on 10 November 1975 relative to depth below mean sea level. - 30 -Table 3 indicates that at both Sites I and 2 more than 90% of the litter collec ted was composed of the seaweeds most characteristic of each area, Nereocystis luetkeana and Laminaria/Agarum for Sites 1 and 2, respectively. The lack of a significant Nereocystis luetkeana contribution to the litter at Site 2 supports the interpretation that litter is not transported long distances away from its place of deposition. The nearest living Nereocystis luetkeana plant to Site 2 was no closer than 100 m. The large accumulation of Laminaria litter at Site 2 may be due to its sheltered location, thereby rendering the area particularly suitable for retention of litter deposited within the immediate vicinity. Within Site 1 there was a similar tendency for litter to be re tained in shelters or pockets formed by the substrate. All large deposits of litter were found in depressions or where the slope of the substrate was more gradual than usual. This can be confirmed by referring to the depth contours for Site 1 (Figure 4). Comparison of the regions of litter retention (Figure 2) to the contour lines demonstrates that the greatest accumulations of litter are where recognizable depressions in the substrate exist. It is important to note that Tridaea cordata and Fucus distichus litter collected in separate poc kets, although the pocket containing Iridaea cordata is only 1.1 m deeper than the pocket containing Fucus distichus. This is further evidence that litter tends to remain in the zone where it was deposited. This effect is less evident in the outer extent of Site 1 where much less litter was collected. Litter entrapment in this region is facilitated by rocks and boulders which provide the topographic relief aiding in the retention of the litter. The seasonal trend in the biomasses of specific and total litter collected within Site 1 is presented in Figure 5 (a-f). The most important fea ture of each of these profiles is that a peak period of litter accumulation occurs in August or September in both of 1975 and 1976, with a low near zero in January and February 1976. Figure 5c demonstrates that the presence of Nereocystis luetkeana stipes in the litter is prolonged over the autumn season. - 31 -Table 3. Comparison of the total quantity and specific composition of litter collected within the transect at 95 m within Site 1 on 9 November 1975 and the total quantity and specific composition of litter collected within the transect at Site 2 on 10 November 1975 (g AFDW/transect)• Site 1 and Site 2 are separated by ca 200 m, the latter being a less exposed area. Species 95 m within Site 1 (j0 Site 2 (%) Fucus distichus 0.66 ( 0.66) 7.81 ( 0.49) Iridaea cordata 0.51 ( 0.51) 16.70 ( 1.05) Nereocystis luetkeana (stipe) 90.28 (90.28) 2.71 (0.17) Nereocystis luetkeana (lamina) 4.79 ( 4.79) 4.09 ( 0.26) Laminaria 1.24 ( 1.24) 1385.26 (86.77) Agarum * 1.82 ( 1.82) 94. 39 ( 5.91) All other species 0.71 ( 0.70) 85.81 ( 5.37) TOTAL 100.01 1596.47 * Agarum fimbriatum Harvey & Agarum cribrosum (Mertens) Bory - 32 -Figure 4. Depth contours (m below mean sea level) for Site 1. Contour intervals are 0.5 m. - 33 -Figure 5. Seasonal distribution of litter biomass for the major contributors to the litter pool within Site 1 based on collections along the 95 m transect location at 3-4 week intervals for the period 20 August 1975 until 2 October 19 76. Contour intervals are g ash-free dry weight per 10 m2. Contour interval a) Fucus distichus 5 .0 b) Iridaea cordata 5 .0 c) Nereocystis luetkeana (stipe) 5 .0 d) Nereocystis luetkeana (lamina) 5 .0 e) Laminaria 5 .0 f) Total litter 10 .0 a) Fucus distichus a c) Nereocystis luetkeana (stipe) 0 230.0 275.0 320.0 1975 365.0 45.0 90.0 DRY OF THE YEAR T 135.0 SI 0 1 N 1 D I JIFIMIfllMIJIJI 1976 ~i 180.0 225.0 270, d) Nereocystis luetkeana (lamina) DRY OF THE YEAR e) Laminaria f) Total litter DRY OF THE YEAR - 40 -This is expected since Nereocystis luetkeana is the most long-lived of the annual plants which contribute significantly to the litter within Site 1. The stipes prevail in the litter longer than the lamina of Nereocystis luet keana as the lamina are more easily detached during rough weather. Structural Composition of Species contributing to Litter: The results for all 10 seaweed species are presented in Table 4. There are considerable differences in the percentages of soluble, moderately resistant and crude fibre components in each species, but it is evident that some species having similar percentages of these components also display taxonomic and/or morphological affinities. Both species of Laminaria have similar percentage compositions of these components as have the stipe and lamina of Nereocystis luetkeana. Iridaea cordata and Gigartina papillata are both particularly low in crude fibre content. Of all the species analysed, Constantinea subulifera has the least percentage of moderately resistant material (29.4%) and the highest per centage of soluble matter (65.6%). It is followed by Fucus distichus in both of these categories, 32.8% and 60.7%, respectively, for moderately resistant and soluble material. Iridaea cordata has both the least percentage of crude fibre (0.86%) and the greatest percentage of moderately resistant material (71.0%). The variability in the percentages of these components among the various species has facilitated the recognition of correlations between the relative amounts of these components in each species and decomposition parameters of these species. These relationships will be discussed in the context of the appropriate experiments. Of particular consequence is the influence of the percentage content of soluble matter on observed rates of oxygen consumption (Experiment 1) and the influence of the percentage crude fibre content on observed rates of particulate matter consumption (Experiment 2) . - 41 -Table 4. The percentages of each of the soluble, moderately resistant and crude fibre components of the significant species within Site 1. Each value is expressed as a percentage of dry weight biomass. Crude fibre glucose refers to the amount of carbohydrate in the crude fibre component expressed as an equivalent amount of glucose. The soluble content and crude fibre components are means of two determinations. Moderately Crude Fibre Component Soluble Resistant Species Component Component Total As glucose Plocamium coccineum war. pacificum 28.1 Rhodomela larix 30.1 Odonthalia floccosa 40.3 Iridaea cordata 28.1 Gigartina papillata 41.0 Constantinea subulifera 65.6 Fucus distichus 60.7 Nereocystis luetkeana (stipe) 41.1 Nereocystis luetkeana (lamina) 44.7 Laminaria saccharina 41.1 Laminaria groenlandica 36.6 Standard error: 59.2 12.70 (3.39) 60.0 9.86 (4.28) 54.7 5.01 (3.44) 71.0 0.86 (0.58) 57.7 1.30 (1.21) 29.4 4.99 (2.26) 32.8 6.48 (1.86) 55.4 3.48 (2.29) 51.6 3.71 (2.27) 52.6 6.30 (3.14) 55.7 7.67 (3.37) — ±0.62 ±0.61 Litter Decomposition Experiments: The results for all 11 litter bag experiments are presented in Figure 6 (a-k). As one litter-bag from the series of litter bags containing Laminaria saccharina was lost, and there being an apparent similarity between the decomposition rates of both species of Laminaria, the data for these two species were combined. Five curve models were applied to each data set with the minimal residual error being the criterion for acceptance, provided the curve maintained a smooth, negative slope. For plots where a logarithmic curve was chosen to represent the data, 2.0% of original dry weight was arbitrarily chosen to repre sent zero percent for graphic purposes, as this curve model approaches the X-axis asymptotically. The five curve models are as follows: 1. Linear: Y = aX + 100.0 2. Quadratic: Y = aX2 + bX + 100.0 3. Logarithmic: lnY = a(lnX) + 100.0 4. Parabolic: Y = (X - a)2/4b; a2/4b = 100.0 5. Hyperbolic: Y = a + (b/(X - c)); a - (b/c) = 100.0 where: 1 X is the independent variable Y is the dependent variable a, b and c are coefficients ln is the natural logarithm Loss of biomass from the litter bags was rapid but the timing and pattern of decomposition was variable among the species. The lamina of Nereo cystis luetkeana decomposed most rapidly, requiring only six days to disappear from the litter bags. The most slowly decomposing species was Fucus distichus, requiring ca 70 days to disappear from the litter bags. Listed in order of de creasing decomposition rates the remaining species are Iridaea cordata (13 days), Laminaria (ca 14 days), Nereocystis luetkeana stipe (ca 18 days), Gigartina papillata (27 days), Rhodomela larix (27 days), Constantinea subulifera (43 days), Odonthalia floccosa (46 days) and Plocamium coccineum var. pacificum (49 days). - 4tJ -Figure 6. Litter decomposition curves (submodels) calculated from data obtained in the litter bag experiments. The curve model (see text), the coefficients (a,b,c) and the coefficient of determination (r^) are given below for each species. Species Model a) Plocamium coccineum var. pacificum P b) Rhodomela larix L c) Odonthalia floccosa P d) Iridaea cordata Q e) Gigartina papillata P f) Constantinea subulifera P g) Fucus distichus LN h) Nereocystis luetkeana (stipe) LN i) Nereocystis luetkeana (lamina) P j) Laminaria LN a b £ r (%) 49.40 6.099 - 99.45 3.720 100.0 - 98.46 45.69 5.220 - 97.52 -0.448 -1.978 100.00 99.90 27.00 1.823 - 99.94 43.42 4.712 - 97.65 -0.059 4.605 - 94.06 -0.210 4.605 - 99.90 6.022 0.907 - 98.56 -0.277 4.605 - 96.80 L: linear Q: quadratic LN: logarithmic P: parabolic H: hyperbolic PERCENTAGE OF ORIGINAL DRY WEIGHT A PERCENTAGE OF ORIGINAL DRY WEIGHT a—• TIME (DAYS) - 54 -Only Iridaea cordata and Rhodomela larix did not subscribe to a decomposition pattern with a decelerating rate of biomass loss. Iridaea cordata was charac terized by an initial lag phase followed by an accelerating rate of biomass loss. Rhodomela larix maintained a linear decomposition rate. Litter Senescence Experiments: The experiments to determine the time for the significant con tributors to the litter to die were not particularly decisive due to the qualitative nature and infrequency of the observations. The results are presen ted in Table 5. At the time these experiments were performed the significance of the contribution by Fucus distichus was underestimated. The estimated time for a seaweed to die was determined for the shaded condition only. Continual deposition of new litter upon existing litter probably means that most litter is at least partially shaded; therefore this condition was accepted as giving a more realistic estimate of the time required for the death of the seaweed to occur. These data were obtained in order that the time taken for seaweed litter to form detritus could be more precisely modelled. As the specific litter components tested demon strated a similarity in their senescence times, six days was accepted as a general estimate for simplicity in modelling. Fucus distichus may have a longer senescence time, but the overall significance of this error is expected to be minor. Nitrogen Content of Decomposing Litter: The nitrogen content of seaweed litter at various stages of decomposition is presented in Table 6 for the 10 species assayed. The most notable feature of these results is that all species except Iridaea cordata demonstrated an increase in the nitrogen:total biomass ratio of material remaining in the litter bags as decomposition proceeded, - 55 -Table 5. Number of days required for unkilled portions of the major con tributors to the litter pool within Site 1 to leave a 1.0 cm mesh litter bag under shaded and exposed conditions. The 'estimated time for senescence' is an estimation of the number of days required for a specific litter component to die once having entered the litter pool. See text for a full explanation. Estimated time Species Exposed Shaded for senescence Iridaea cordata 24-30 10-14 5 Nereocystis luetkeana (stipe) 24-30 15-2 3 9 Nereocystis luetkeana (lamina) 15-23 6-10 6 Laminaria saccharina 24-30 10-14 5 Laminaria groenlandica 24-30 10-14 5 - 56 -Table 6. Percentage nitrogen content of the material remaining within the litter bags at the termination of their incubation period. Percentage of Species original dry weight Percentage nitrogen Plocamium coccineum var. pad fi cum 100.00 3.74 65.26 3.68 42.50 3.89 28.22 4.64 Rhodomela larix 100.00 4.24 86.20 4.43 48.73 4.74 Odonthalia floccosa 100.00 4.24 55.35 3.71 34.51 4.50 Iridaea cordata 100.00 1.94 97.39 1.955.66 1.63 Gigartina papillata 100.00 2.54 38.50 3.116.79 4.26 2.72 6.34 Constantinea subulifera 100.00 2.61 62.30 2.70 45.20 - 3.17 Fucus distichus 100.00 1.73 61.08 2.21 39.99 2.37 Nereocystis luetkeana (stipe) 100.00 1.50 29.70 2.16 Nereocystis luetkeana (lamina) 100.00 2.38 52.80 3.76 Laminaria saccharina 100.00 1.98 13.70 3.70 Laminaria groenlandica 100.00 2.64 30.14 4.10 11.16 5.2- 57 -although total nitrogen content decreased. The greatest percentage nitrogen content was observed for Gigartina papillata, most likely because it was the most fully decomposed of all the species when final nitrogen content was analyzed. The nitrogen:total biomass ratio of nearly fully decomposed Gigartina papillata increased over that of undecomposed Gigartina .papillata by 250%. Figure 7, which incorporates data from all species assayed, demonstrates that a hyperbolic curve approximates the trend of increasing nitrogen:total biomass ratio very well, indicating an accelerating increase in litter nitrogen content relative to other biomass components as decom position proceeds. Detritus Decomposition: Both experiments tested the following three major effects for their impact on decomposition rates:. 1. Length of the incubation period 2. Source of the detritus, i. e. the seaweed from which it was created 3. Size of the detrital particles For brevity each of these effects will often be referred to as the 'incuba tion period', 'detrital species', and 'particle size' effects, respectively. Experiment 1 (Microbial Oxygen Consumption): An Analysis of Variance (ANOVA) was performed on the oxygen consumption data obtained in this experiment. The results of the analysis are presented in Table 7. Referring to the three major effects, it can be concluded that only two of them, the detrital species and the length of the incubation period, are significant (p < .05) contributors to the observed differences in the oxygen comsumption rates. In consideration of the latter effect, such a response must be expected since the oxygen within gure 7. Plot demonstrating an increase in the ratio of nitrogen:dry weight biomass of decomposing litter expressed relative to a ratio of 1:1 for undecomposed litter. All 10 species assayed are incorporated within the plot. The solid line indicates the best fit through the points. Table 7. Analysis of variance table for the results of Experiment 1, demonstrating the effects of particle size, detrital species and length of incubation period on the oxygen consumption by microbes utilizing the detritus as a carbon source. Source of variance Degrees of freedom Particle size (PS): Detrital species (DS): PS - DS interaction: Incubation period (IP) PS - IP interaction: DS - IP interaction: Residual error: Total: 2 10 20 3 6 30 60 131 Sum of squares 0.12379E-02 0.65865 0.18062E-01 5.1409 0.21561E-02 0.29697 0.82011E-01 6.2000 Mean sum of squares 0.61894E-03 0.65865E-01 0.90311E-03 1.1736 0.35934E-03 0.98989E-02 0.13668E-02 Probability 0.6 3799 0.0 * 0.84772 0.17613E-52 0.95198 0.72414E-10 * significant for a = 0.05 - 60 -the BOD bottle is continually being consumed. That detritus of different biogenic origins contributed significantly to the observed variation in the oxygen consumption rates implies that some species of detritus are more susceptible to breakdown by microbes than others. For the third major effect, particle size, there was no detectable difference among the oxygen consumptions of the three particle sizes. Any response that may have occurred could have been easily attributed to chance. The second source of significant variation within the experiment can be explained in terms of an interaction between the detrital species and their response over the incubation periods. The essence of the interaction is that utilization of the oxygen in the BOD bottles follows a pattern dependent upon the biogenic origin of the detritus. By observing Figure 8, which relates the cumulative oxygen consumption to the length of the incubation period for all 10 species, it can be seen that the significance of the interaction term is a result of the relatively steep slope maintained by Fucus distichus during the 10-20 day incubation period and to the heterogeneity of the slopes within the 10-15 day incubation period. Three 'a posteriori' range tests were performed on the data in an attempt to delimit affinities and detect outliers among the responses to the significant major effects. These tests were: 1. Duncan's New Multiple Range Test 2. Newman - Keul's Test 3. Tukey's Test Not unexpectedly, each incubation period (0,5,10, and 20 days) was rendered unique and independent. Only Duncan's Test defined exclusive subsets for the effect of detrital species on oxygen consumption rates. Both the Newman - Keul's Test and Tukey's Test permitted entities to have 61 -Figure 8. Cumulative oxygen consumption by microbes decomposing the 10 detrital species in Experiment 1. Each data point is the mean result for the three detrital particle sizes. Plocamium coccineum var. pacificum Rhodomela larix Odonthalia floccosa Iridaea cordata Gigartina papillata Constantinea subulifera Fucus distichus Nereocystis luetkeana (stipe) Nereocystis luetkeana (lamina) Laminaria saccaharina Laminaria groenlandica - Ua-0.0 5.D 10.0 15.0 20.0 INCUBATION TIME (DAYS) - 62 -a membership in more than one subset such that affinities were more diffi cult to detect. The subsets defined by Duncan's Test are presented in Table 8a. To test for the possible influence of the soluble compon ent on the results obtained, the oxygen consumed by each detrital species after five days of incubation (mean of three particle sizes) was regressed on the percentage soluble content of each species. The result is signifi cant (p < .01) and conclusive if Iridaea cordata detritus is excluded from consideration. The relationship between oxygen consumption and soluble content is presented in Figure 9. About 77% of the variation in oxygen consumption can be accounted for by differences in the soluble content of the detrital species. Reference to Table 8b indicates the mean percentage soluble content of the species comprising each subset. The trend of increasingly higher percentage soluble contents for the subsets characterized by the more rapidly decomposing species is evident, with the exception that Iridaea cordata decomposes rapidly although containing a relatively small percentage of soluble matter. Experiment 2 (Microbial Consumption of Particulate Material): The results of an ANOVA on the decomposition data obtained in this experiment are presented in Table 9. Reference to it shows that there are four significant sources of variation (p < .05) . As was the case in Experiment 1, only two major effects were significant, incubation period and detrital species. Two other significant sources of variation were an interaction between incubation period and detrital species and an interaction between particle size and detrital species. Table 8. a) Subsets delimited by Duncan's New Multiple Range Test. Each subset contains those detrital species which show a significant (p < .05) degree of affinity with respect to the quantity of oxygen consumed by microbes decomposing the detritus. Subset 1 Subset 2 Subset 3 Subset 4 Plocamium coccineum va r. pacificum Gigartina papillata Rhodomela larix Odonthalia floccosa Nereocystis luetkeana (stipe) Nereocystis luetkeana (lamina) Laminaria saccharina Laminaria groenlandica Iridaea cordata Constantinea subulifera Fucus distichus b) The average percentage soluble content of the subsets in Table 8a. Subset 1 Subset 2 Subset 3 Subset 4 34.9 ± 7.2% 40.9 ± 4.1% C. subulifera 65.6 ± 2.8% I. cordata 28.1 ± 0.1% 60.7 ± 3.8% Figure 9. Relationship between the percentage soluble contents of the 10 detrital species (exclusive of Iridaea cordata) and the quantity of oxygen consumed by microbes decomposing the detritus after five days of incubation, as determined in Experiment 1. The solid line indicates the best fit through the points. Table 9. Analysis of variance table for the results of Experiment 2, demonstrating the effects of particle size, detrital species and length of incubation period on the consumption of particulate material by microbes utilizing detritus as a carbon Source. Source of variance Degrees of freedom Particle size (PS) : Detrital species (DS): PS - DS interaction: Incubation period (IP) PS - IP interaction: DS - IP interaction: Residual error: Total: 2 10 20 3 6 30 60 131 Sum of squares 135.22 41877.0 1875.8 14200.0 363.59 15227.0 3116.1 76795.0 Mean sum of squares 67.611 4187.7 9 3.788 4733.4 60.599 507.56 51.9 35 Probability 0.27960 0.0 * 0.4082 3E-01 0.95989E-21 0.33608 0.15329E-12 * significant for a = 0.05 - 66 -The significant response for the length of the incubation period was expected as the general trend would be a continual loss of particulate mater ial as time proceeds; however, this did not always occur. The second signifi cant response was due to the different decomposition rates of the various detrital species. Figure 10 indicates an initial decay rate for Iridaea cordata which would reduce it to zero in 18 days. In comparison, there appear to be some anomalous results for Plocamium coccineum var. pacificum, Rhodomela larix, Odonthalia flocossa, and Fucus distichus, all of which show an increase in dry weight of particulate matter following 10 days of incubation. As there was a significant interaction between detrital species and incubation period, range tests were performed to delimit any groupings which might provide insight into the reasons for the interaction. None of the three range tests delimited exclusive subsets. The most definitive was Newman -Keul's Test which delimited five subsets, with Laminaria groenlandica being a member of two of them. As the overall mean for Laminaria groenlandica was closer to that of Laminaria saccharina than of Nereocystis luetkeana (stipe), its near est neighbours in each of the subsets in which it was placed, it was placed with Laminaria saccharina. This rendered all the subsets unique in composition. The composition of the subsets is presented in Table 10. There is also a significant interaction between particle size and detrital species. The implication is that there may be some species of detritus whose decomposition rate is dependent upon the size of the detrital particles. Closer inspection of the data revealed that the two most rapidly decomposing spe cies , Iridaea cordata and Nereocystis luetkeana (stipe and lamina sections com bined) , displayed a trend toward a more rapid decomposition rate as mean parti cle size decreased. A detectable difference in decomposition rates in response to particle size is most likely for the most rapidly decomposing species since the experimental error would be a smaller proportion of the total variance than - 67 -Figure 10. Cumulative loss of particulate material from the 10 detrital species decomposed in Experiment 2. Each data point is the mean result for the three detrital particle sizes. Plocamium coccineum var. pacificum Rhodomela larix Odonthalia floccosa Iridaea cordata Gigartina papillata Constantinea subulifera Fucus distichus Nereocystis luetkeana (stipe) Nereocystis luetkeana (lamina) Laminaria saccharina Laminaria groenlandica PERCENTAGE OF ORIGINAL DRY WEIGHT Table 10. Subsets delimited by Newman -detrital species which show a significant (p quantity of particulate material consumed by Keul's Range Test. Each subset contains those < .05) degree of affinity with respect to the microbes decomposing the detritus. Subset 1 Subset 2 Subset 3 Subset 4 Subset 5 i Plocamium coccineum Gigartina papillata Laminaria saccharina Nereocystis luetkeana Iridaea cordata var. pacificum (stipe) co Laminaria groenlandica 1 Fucus distichus Nereocystis luetkeana (lamina) Rhodolema larix Odonthalia floccosa Constantinea subulifera - 69 -for less rapidly decomposing species. Figure 11 (a,b) graphically presents the results for both of these species. Note in particular that the difference in decomposition rates for the three particle sizes is most evident after only 10 days of incubation, while the conditions within the culture vessels are still sufficiently fresh to maximize the experimental effects. Because of adverse effects caused by the lengthier periods of incubation, the effect of particle size could not be shown to be statistically significant for either species. Two regression analyses were performed to test the hypothesis that the decomposition rates of seaweed detritus were at least partially a func tion of the crude fibre content of the detritus. The dependent variable in both cases was the maximum percentage loss (mean of 3 particle sizes) of particulate ma terial observed for each detrital species in Experiment 2. For the species which showed an initial increase in dry weight as time proceeded, the rate of loss of particulate material was determined in relation to the maximum dry-weight attained. The independent variables were crude fibre content and crude fibre carbohydrate expressed in glucose equivalents. Both were expressed as a percentage of the total particulate component (crude fibre plus moderately resis tant material). The results of both regression analyses were significant(p < .05). Figure 12a demonstrates the relationship between maximum percentage loss of par ticulate material and percentage crude fibre content. Figure 12b presents an equivalent relationship using percent glucose as the independent variable. Since decomposition rates would theoretically never be expected to reach zero, an exponential decay curve was considered the most appropriate model. The re gression analyses accounted for 42.8% and 39.9% of the variance observed in Figures 12a and 12b, respectively. - 70 -Figure 11. Cumulative loss of particulate material from a) Iridaea cordata b) Nereocystis luetkeana (stipe and lamina combined) detritus. For each species the results for the three detrital particle sizes are presented. The three par ticle sizes are as follows: a) 1000-420 urn 0) 250-149 urn y) 44-0 ym PERCENTAGE OF ORIGINAL DRY WEIGHT _T>0£-- 71 -Figure 12. Relationship between the maximum percentage loss of particulate material from the 10 detrital species decom posed in Experiment 2 and the percentage of crude fibre in the particulate material of each detrital species. The solid lines indicate the best fit through the points. a) crude fibre expressed as a percentage of the dry weight of the particulate material. b) crude fibre carbohydrate expressed as an equivalent amount of glucose and as a percentage of the dry weight of the particulate material. PERCENTAGE OF CRUDE FIBRE GLUCOSE - 72 -Detritus Assessment: The biomass of detritus along the permanent transect location within Site 1 is best represented graphically by Figure 13. This three-dimen sional representation demonstrates that the availability of detritus reached a maximum of ca 1.4 g AFDW/m about the middle of August in 1976. The peak occurs near the time of maximum litter biomass and within the central zone of the sea weed bed. The quantity of detritus diminishes towards the inner and outer edges of the bed to 15-30% of the maximum value. During the summer of 19 75 natural detritus was periodically ex amined microscopically for characteristics which might aid in determination of its origin. Its composition was determined to be amorphous, consisting mostly of variously shaped colourless unidentifiable particles, as well as some dia-tomaceous material. The latter accounted for ca 10% of the material observed. Faunal Assessment: In order to recognize a coincidence of the occurrence of specific fauna and the maximum availability of detritus» the sums, by numbers and dry weights, for each species occurring within the summer faunal collections of 28 July, 18 August and 12 September were expressed as a percentage of the total for the seven collections from May until October. These results are presented in Tables 11a (numbers) and lib (dry weight). Those species whose occurrence coin cides with the maximum availability of detritus were delimited on the basis of the following, somewhat arbitrary, criterion. The qualifying species must have been represented by more than 75% of their total number and dry weight during the summer collections. The three species which met this qualification were: Cancer oregonensis Dall Wetacaprella. anomala Mayer Lacuna marmorata Dall - 73 -Figure 13. Contour representation of detritus biomass along the 95 m transect location within Site 1 for the period 28 May until 7 October 1976. Contour intervals are 0.2 g ash-free dry weight per nr. - 74 -Table 11a. The total number of each faunal species summed over the 28 July, 18 August and 12 September 1976 transect collections. The percentage that this number represents of the total number of occurrences over the entire sampling period is in parentheses. An * denotes those species which are represented by more than 75% of their total number of occurrences within the samples collected on the above three dates. Species Number Percentage of total Acmaea mitra Rathke 1 (12.5) Alvinia spp. 119 (16.9) Amphilochus sp. 11 (44.0) Amphithoe sp. 1 (1.4) Balcis mi cans Carpenter 9 (60.0.) Bittium eschrichtii Middendorff 56 (49.1) Cancer oregonensis 19 (100.0) * Chlamys hastatus Sowerby 4 (33.3) Clinocardium sp. 12 (34.4) Granulina margaritula 161 (54.9) Hemigrapsus nudus Dana 3 (50.0) Hiatella arctica L. 6 (31.6) Lacuna marmorata 6018 (88.6) * Lirularia lirulata 66 (57.4) Margarites pupillus Gould (juvenile) 1111 (37.6) Margarites pupillus (parental) 449 (39.5) Metacaprella anomala 9 (100.0) * Mitrella gouldii Carpenter 109 (46.5) Mytilus edulis L. 858 (30.6) Nereis pelagica L. 1 (11.1) Notoacmea scutum Rathke 8 (40.0) Ocenebra sp. 25 (48.1) Odostomia spp. 116 (70.1) Pag.urus kennerlyi Stimpson 1 (8.3) Pugettia richii Dana 10 (30.4) Strongylocentrotus droebachiensis 4 (33.3) Tonicella lineata Wood 53 (31.8) - 75 -Table lib. The total dry weight of each faunal species summed over the 28 July, 18 August and 12 September 1976 transect collections. The percentage that this figure represents of the total dry weight of individuals collected over the entire sampling period is in parentheses. An * denotes those species which are represented by more than 75% of their total dry weight within the samples collected on the above three dates. Species Dry Weight Percentage of total Acmaea mitra 2.178 (14.8' Alvinia spp. 0.1963 (18.6) Amphilochus sp. 0.0100 (33.6 Amphithoe sp. 0.0029 (0.7] Balcis mi cans 0.0378 (57.7 Bittium eschrichtii 3.474 (42.1 Cancer oregonensis 0.1142 (100.0) Chlamys hastatus 0.0460 (41.0 Clinocardium sp. 0.3748 (31.6 Granulina margaritula 0.4882 (54. 3) Hemigrapsus nudus 1.115 (94.2) Hiatella arctica 1.66 7 (47.1, Lacuna marmorata 12.2 3 (75.0 Lirularia lirulata 0.6519 (52.1 Margarites pupillus (juvenile) 3.247 (26.9] Margarites pupillus (parental) 11.49 (42.7) Metacaprella anomala 0.0108 (100.0) Mitrella gouldii 3.711 (41.5) Mytilus edulis 62.94 (62.1) Nereis pelagica 0.0079 (4.5) Notoacmea scutum 2.163 (42.7) Ocenebra sp. 1.061 (21.5) Odostomia spp. 0.3368 (63.3) Pagurus kennerlyi 0.1475 (55.1) Pugettia richii 1.942 (27.9) Strongylocentrotus droebachiensis 0.4500 (6.3) Tonicella lineata 8.064 (21.0) - 76 -For each of these species histograms are presented in Figure 14 (a-c) to des cribe the temporal distributions of numbers and dry weight over the period sam pled, permitting a graphic interpretation of their seasonal abundances. All three species demonstrate a trend of increasing numbers and biomass toward a strong midsummer peak followed by a decrease in these parameters in September and October, implying that the sampling program is a sufficient documentation of their seasonal abundance patterns in 1976. For Lacuna marmorata, which was particularly abundant throughout the summer months, additional trends were evident. Concomitant with an increase in numbers and dry weight of Lacuna marmorata is a decrease in the mean dry weight per individual. Figure 15 indicates that the greater increase in numbers relative to dry weight appears following the second sampling date (14 June 1976) and is due to the occurrence of a large number of juvenile individuals. Most Lacuna marmorata, and in particular the juveniles, were generally found amongst the detritus and debris accumulated on the bottom and consolidated by the plants comprising the subtidal turf community. There is evidence that the abundance of juvenile Lacuna marmorata in the detritus and debris is due to their utilizing the detritus as a food re source. Figure 16 demonstrates that 100% of the total number and dry weight of Lacuna marmorata were collected within the quadrats at 30, 40 and 50 m along the transect. Results from the detritus collections of 20 August 1976 determined this to be the area where most detritus retention occurred. - 77 -Figure 14. Seasonal distribution histograms of the total number and dry weight (g) of a) Cancer oregonensis b) Metacaprella anomala c) Lacuna marmorata occuring within the seven transect collections from 25 May until 7 October 1976. Open bars: numbers Solid bars: biomass NUMBER 0 2.5E3 5.0E3 I I I - 78 -Figure 15. Seasonal trend in the mean dry weight (g) per individual of Lacuna marmorata for the period 25 May until 7 October 1976. The occurrence of juvenile individuals is evidenced by a decrease in the mean dry weight per individual after the second (14 June) sampling date. rv a Di <C Q-UJ a JS CS s in CN 3 co 3 CO CN -P tn 2 < co u CD -P OH Q) 1/3 M <D Xi o +J o o - 79 -Figure 16. o CD s: \ r-i CD *—' CXI 1— T-H m CD 00 LU d >-cr: d o o o 00 LU o 1 CD Q_ O CO 1 <n o 1— CM o I— 11 O O i i i O CD 00 <C h— O LU CD C_> cr: LU O Q_ o CSl o Spatial distribution along the 95 m transect location within Site 1 of Lacuna marmorata (numbers and biomass) and detritus biomass demonstrating a coincidence in the occurrence of their maximum abundances. * A collection at 20 m on this date contained no Lacuna marmorata. Detritus biomass (20 August 1976) Lacuna marmorata (numbers) (18 August 1976) Lacuna marmorata (biomass) (18 August 1976) 20.0 30.0 40.0 50.0 60.0 70.0 80.0 90.0 100.0 DISTANCE ALONG TRANSECT (M) DISCUSSION Litter Assessment: In order to assess detritus formation and subsequent processing within Site 1, it is necessary to know the contributions of significant sources of detritus within the system. In such a coastal area there is potential for input from both terrestrial and marine sources. Bath Island is removed from salt marshes and domestic sewage input, so the potential sources are reduced to plankton, faunal excreta, drift wood and seaweed. Plankton will not be an im portant contributor, at a biomass ca 1% that of seaweed (Blinks 1955). Faunal excreta is not likely to exceed that amount as it is two trophic levels removed from plant production. Perhaps the most significant allochthonous litter source in British Columbia waters is drift wood (Perkins 1974). The seaweed zone of Bath Island is not characterized by a noticeable settlement of wood particles, such that seaweeds can be considered the only important source of litter. It is difficult to compare litter accumulation at Bath Island to other areas as quantities are a function of the geology and biology of the area being studied. Zobell (1971), in a study of drift seaweeds cast upon San Diego County beaches in California between 19 36 and 1954, estimated as much as 184 of seaweeds per 150 m of shoreline on the beach at certain times. In contrast to Zobell's results virtually no litter was collected intertidally or supratidally at Bath Island, and the quantity of litter collected subtidally was very small in comparison to Zobell's intertidal assessment. The difference in the regions of litter deposition is attributable to beaches being accretion areas whereas rocky shores are excretion areas. The only significant comparison that can be made is that the phaeophytes were the dominant contributors to the litter pool in both studies, at 75% and 86% for California and Bath Island, respectively, on a wet weight basis. - 81 -The seaweeds contributing the most biomass to the litter in Site 1 are relatively productive species. Both Jridaea cordata and Nereocystis luetkeana grow rapidly during the spring, attaining their maximum standing crop biomasses during the summer. This is followed by a period of increasing litter deposition. At this time the plants have reached a size where they become less able to withstand the onslaught of current and waves, their vulnerability resul ting in some plants becoming detached from the subtrate. For Nereocystis luet keana the lamina are the more significant contributors to the litter. Occasion ally healthy plants become detached at their bases and drift subject to the effects of winds and current, due to a pneumatocyst keeping the plants afloat. Neither the fate of these plants nor the number that left Site 1 during the course of this study are known; however it is known that they are not generally cast ashore at either Site 1 or Site 2. Rarely was a Nereocystis luetkeana litter fragment observed upon the shore and only once during the entire study was Nereocystis luetkeana litter collected within an intertidal quadrat. This is not unexpected as rocky shores are areas of excretion. Laminaria and Fucus dis tichus, although tending to be perennial, contribute significantly to the litter pool after having finished most of their seasonal growth. Slower growing seaweeds are disproportionately represented in the litter collections. Constantinea subulifera, a dominant, long-lived contributor to seaweed standing crop biomass in Site 1 was not collected during the litter sampling program. By virtue of the sampling scheme undertaken, it has been possible to assess the biomass of seaweed litter within Site 1 in four dimensions. The midsummer collections create an areal profile for the site and delimit the spatial * characteristics of litter distribution. The 14-month sampling program at the 95 m transect location within Site 1 contributes a temporal dimension. This - 82 -facilitates an extrapolation of the midsummer areal profile over the period of a full year for the five species contributing significantly to the litter. It will be seen in a later discussion concerning the development of a mathematical model to simulate litter decomposition that the most impor tant observation with respect to the temporal distribution of litter biomass is the similarity of the seasonal patterns of the five major contributors. Although the longevity of Nereocystis luetkeana stipes within the litter exceeds that of other litter, their total contribution is relatively minor. Any loss of informa tion resulting from usage of a single curve model to approximate the seasonal distribution for total litter will be almost negligible. The quantity of litter available for decomposition is the ulti mate driving variable in an attempt to simulate its entry into, and processing within, litter and detrital pathways. Although these collections are most repre sentative of the litter distribution patterns within Site 1, one must be careful not to accept immediately that these data represent the true proportion of each species' contribution to total litter input since neither the decomposition rate for each species nor the residence time for litter in Site 1 have been considered. Litter Decomposition Experiments: There are three components of plant litter which are known to in fluence its decomposition rate. The soluble, moderately resistant and crude fibre components respond differently during the decomposition process. Extrinsic influences must be considered as well. Environmental factors such as temperature, moisture, nutrient availability and microbial composition interactively exert an effect on the decomposition rates and patterns adding to the complexity of the process. Some authors (Grill and Richards 1964, Minderman 1968, Otsuki and Hanya 1972) chose a 'theoretically preferable' curve model to represent their data on litter decomposition which Olson (1963) introduced as an exponential - 83 -decay curve with a constant 'k' related to the half-life of the substance under going decomposition. For such a simple model to be satisfactory the process it describes must be simple as well. This is not the case with litter decomposi tion. Although the curve may adequately describe the individual components of the decomposition process, their combination may defy a simple description. Hunt (1977) demonstrated this point well and illustrated the unsuitability of applying an exponential decay curve to the decomposition data of Pendleton (1972). The result produced curves which obviously misrepresented the data. With this in mind it was more suitable to select a curve which extrinsically fit the data well rather than accept an intrinsic model based solely on theoretical considerations. Furthermore, acceptance of an intrinsic model re duces the probability for success of a practical application of such information when compared to efforts based on a more realistic representation of the data. The most consistent trend observed in the litter bag experiments was the initial rapid loss of material followed by a decrease in this rate as time proceeds. A similar trend is normally observed for terrestrial litter, al though over a much longer time period, and is explained as follows. There is an initial rapid loss through leaching of the soluble and more easily metabolised components (Nykvist 1963, Petersen and Cummins 1974, Suberkropp et al. 1976) leaving behind a structural backbone of refractory material which slowly decompo ses over a period of months (Lousier and Parkinson 1975, Stachurski and Zimka 1975, 1976 a&b, Gasith and Lawacz 1976). As the refractory material becomes more prev alent its resistance to metabolism by microbes results in the decomposition pro cess slowing down. Only two species, Rhodomela larix and Iridaea cordata, differed from this trend. Iridaea cordata displayed an accelerating decomposition rate as time proceeded while Rhodomela larix maintained a linear rate of decomposition. Iridaea cordata is unique by having the lowest observed percentages of soluble and crude fibre components. Having a low soluble content would reduce the • - 84 -length of an initial period of leaching and the paucity of crude fibre would facilitate a relatively rapid decomposition rate following this period. The initial lag phas'e may be due to the maintenance of structural integrity during the primary stages of decomposition and the inability of the small amount of soluble matter to mask this effect as it apparently does for other species. The linear decomposition curve for Rhodomela larix can perhaps be explained by its having a predominance of short, stubby branches which may become suitably fractured to escape the litter bag before its relatively high crude fibre content limits its decomposition rate in the later stages. Loss of seaweed biomass from the litter bags was rapid. When compared to terrestrial litter, seaweed litter decomposes at least five times faster. Odum and de la Cruz (1967) and de la Cruz (1975) demonstrated that salt marsh plants decompose at about the same rate as terrestrial plants. Most plants they studied had a considerable amount of their original dry weight remaining after 300 days. Similarly, de la Cruz and Gabriel (1974) determined loss of Juncus roemerianus Scheele from litter bags to be ca 40% per year. Adding aquatic vascular plants to the comparison, Harrison and Mann (1975b) found that Zostera marina lost only 35% of its original dry weight in 100 days, under laboratory conditions. Hunter (1976) studied two freshwater plants, Lemna minor L. and Chara contraria A. Braun ex Kutzing, and found both to re tain ca 75% of their original dry weight after ten weeks of submersed incubation in litter bags. That terrestrial, aquatic and marine vascular plants decompose much more slowly than seaweeds, even when submersed, implies that the rapidity of seaweed decomposition is more a function of their composition than of their environment. The influence of the relative quantities of seaweed structural components on decomposition rates is discussed in relation to the detritus de-: composition experiments (Experiments 1 and 2). - 85 -Nitrogen Content of Decomposing Litter: The results of this study are particularly significant in that they demonstrate a difference between vascular plant decomposition and seaweed decomposition with respect to nitrogen content. For vascular plant litter there is generally an increase in both the concentration and absolute content of nitro gen following an initial period of leaching during which most of the soluble com ponents escape (Nykvist 1963, Petersen and Cummins 1974, Suberkropp et al. 1976). As most nitrogen escapes as soluble matter it has to be reacquired from the sur rounding environment by organisms associated with the litter (Bocock 1964). Al ternatively, this study indicates that the relative increase in nitrogen content of decomposing seaweed litter is due to a preferential release of chemical con stituents low in nitrogen. That the increase is due to the incorporation of inorganic nitrogen into the litter by the activity of microbes is unlikely as it requires that the microbes have phenomenally rapid growth and nitrogen incorpora tion rates at a time when inorganic nitrogen in the seawater at Site 1 is at a low concentration (Tully and Dodimead 1957). This argument is enhanced by demonstrating that C:N ratios of 8:1 or less would be very difficult to attain by metabolic processes. C:N ratios of this order are implied by the data obtained in this study. If the highest percentage nitrogen contents obtained for Laminaria saccharina (3.70%) and Rhodomela larix (4.74%) are related to the percentage carbon contents for Lamin aria saccharina (26.76%) and an unspecified Rhodomela (28.32%) (Vinogradov 1953), a C:N ratio of 6-8:1 results. With a value of 6.4% nitrogen content obtained in this study for 9 7% decomposed Gigartina papillata and a value of 24% carbon con tent for Gigartina acicularis (Wulfen) Lamouroux (Niell 1976) a C:N ratio of less than 4:1 results, assuming there is a reasonable degree of generic similarity in percentage carbon contents. The C:N ratio for bacteria is ca 5.7:1 (Spector 1956). To attain - 86 -C:N ratios approaching this figure the material in the litter bags would have to be composed almost entirely of microbial biomass, unless a considerable propor tion of the nitrogen was a component of the initial seaweed biomass. Whyte and Englar (1975) suggest that a large proportion of the protein in seaweeds, Nereo cystis luetkeana in particular, is bonded to the cellulosic fibres of the cell wall. This would prevent the easy release of protein nitrogen since cellulose is a particularly resistant component. The hyperbolic curve presented in Figure 7 is consistent with a protein - cellulose bond hypothesis. An accelerating in crease in relative nitrogen content implies the rate of nitrogen loss is inde pendent of the rate of loss of the more abundant biomass components. Hunter (1976) used litter bags to assess the decomposition rate of Fucus vesiculosus on a rocky shore and within a salt marsh. His results are comparable to those presented in this study with respect to decomposition rates, relative nitrogen content and C:N ratio. Additionally, for the aquatic plants Lemna minor and Chara contraria, he demonstrated no consistent trend for the same parameters, maintaining the uniqueness of seaweeds in this regard. Detritus Decomposition: For the subsets delimited in Experiment 1 (Table 8a) the within-group affinities are somewhat apparent. All groups are composed of species of a single taxonomic class and can be categorized according to the morphology and habit of the seaweeds they contain. Subset 1 contains four species of branched Rhodophyta which are found intertidally or in the shallow subtidal zone. Sub set 2 contains three species of subtidal kelp (Laminariales). One other phae-ophyte, Fucus distichus, is placed by itself in Subset 4. It resembles the other phaeophytes neither in morphology nor habit, being dichotomously branched and inhabiting the intertidal zone. Subset 3 contains two bladed rhodophytes known to coexist in the shallow subtidal zone (Foreman unpub.). Similarly, for Experiment 2, the within group affinities can be easily detected. Subset 1 contains all the 'resistant' species, those which did - 87 -not exhibit a continual • los.s of particulate biomass, and Constantinea subuli fera. Constantinea subulifera decomposed faster than all other species in Subset 1 and is the only species that did not exhibit an increase in particu late biomass as decomposition proceeded. It is the only bladed seaweed in Subset 1. Although Newman-Keul's Test did not separate Constantinea sublifera from the other species, Duncan's Test delimited an equivalent subset excepting Constantinea subulifera. The increase in particulate biomass can be explained as a result of increased microbial biomass due to preferential metabolism of soluble matter as demonstrated in Experiment 1. If the growth rate of microbes utilizing the soluble matter exceeds the decomposition rate of the particulate fraction of the detritus a net increase in particulate biomass will result. Fucus distichus (60.7%) and Constantinea subulifera (65.6%) have soluble con tents considerably higher than the eight other species. Although low in soluble content, Rhodomela larix (4.28%), Odonthalia flocossa (3.44%) and Plocamium coccineum var. pacificum (3.39%) have the highest percentages of crude fibre carbohydrate. Experiments 1 and 2, respectively, demonstrated that soluble matter is preferentially metabolized and decomposition is slower for seaweeds with a high crude fibre content. Laminaria saccharina and Laminaria groenlandica comprise Sub set 3. Subset 4 contains the lamina and stipe sections of Nereocystis luet keana. Kelp being delimited from the other seaweeds indicates taxonomic similarities with regard to decomposition susceptability. Subsets 2 and 5 contain single species each. Iridaea cordata is isolated because of its rapid decomposition rate. Gigartina papillata is placed between Subset 1 and the subsets containing more easily decomposable species. It is the only intertidal species not contained within Subset 1. It has an affinity with the species in the other three subsets, all of which are bladed seaweeds, in that it has a tendency to be foliose. - 88 -There is a basic similarity in the composition of the subsets delimited in Experiment 1 and Experiment 2. Any differences can readily be ex plained by the influence of detrital soluble matter content on the rates of oxy gen consumption obtained in Experiment 1. Referring to Table 8a, the effect of the high soluble contents of Fucus distichus and Constantinea subulifera on their oxygen consumption rates can be negated by placing them in Subset 1 with the other 'resistant' species. All four subsets are now less dissected equi valents of those delimited in Table 10. The inability to dissociate Nereocys tis luetkeana from Laminaria saccharina and Laminaria groenlandica, and Gigar tina papillata from the other members of the Subset 1 is likely due to the an alysis becoming less powerful as a result of the error contributed by the co-variance of oxygen consumption with the quantity of soluble matter in the detri tus, as demonstrated by Figure 9 and Table 8b. As a final judgement, three generalizations can be made concern ing the decomposition rates observed. Detritus derived from intertidal seaweeds is apparently more resistant to decomposition than detritus derived from sub tidal seaweeds. Detritus derived from the faster growing seaweeds decomposes more quickly than detritus derived from the slower growing seaweeds. Seaweed morphology appears to correlate with decomposition susceptibility, the more foliose the seaweed, the more quickly detritus derived from the seaweed decomposes. All three of the above considerations are closely interrelated. Other factors are likely to be involved as well, in particular the resistance of seaweeds to at tack by microbes. The presence of antibacterial chemicals in some species is known to enhance resistance (Sieburth 1968). In Experiment 1 oxygen consumption rates were shown to correlate with the soluble content of specific detritus (Figure 9). Consumption was higher for species having relatively high soluble matter contents. Only Iridaea cor data defied this trend. The oxygen consumption rate of microbes decomposing - 89 -.Iridaea cordata detritus was second only to Fucus distichus, although it has the lowest percentage soluble content (28.1%) observed amongst all species. Its rapid decomposition rate may be partially due to it containing only a very small quantity of crude fibre at 0.86% of its dry weight. This was the lowest quantity observed amongst all the species. This lack of resistant material may render Iridaea cordata more vulnerable to attack by microorganisms such that it decomposes rapidly relative to other species with a similar or greater percentage of soluble matter. The decomposition rates obtained in Experiments 1 and 2 are com plementary. Iridaea cordata provides the best comparison due to its having very little crude fibre, a low soluble matter content, and a rapid decomposition rate. By assuming that the particulate component of detritus is composed mostly of carbohydrate, ca 1.07 mg of oxygen would be required to fully decompose the 1.0 mg plug of detritus introduced into each BOD bottle. At the average oxygen consumption rate of 0.052 mg O2 per day observed for Iridaea cordata over the first 10 days of incubation, 20.5 days would be required to fully decompose the detritus. The rate of loss of particulate matter obtained for Iridaea cordata during the first 10 days of incubation in Experiment 2 was 5.7% per day. At this rate, 18 days would be required to fully decompose the detritus, in close agree ment with the 20.5 days estimated by the oxygen consumption method. Comparing the decomposition rates obtained in Experiments 1 and 2 to those obtained by other persons for vascular plant detritus, the most ap parent difference is the relative rapidity of seaweed detritus decomposition. Odum and de la Cruz (1967) measured oxygen consumption of natural coarse Spar-tina alterniflora detritus (that which was retained by a 0.239 mm aperture) at ca 1.8 mg 02/g AFDW/hr at 15 C. This study obtained rates in the range of .2.7-7.0 mg 02/g AFDW/hr for equivalently sized detritus from Plocamium coccineum var. pacificum and Iridaea cordata, respectively, also at 15 C. The data were - 90 -most reliable for these two species since their low soluble contents minimally affected observed oxygen consumption rates. Differences in decomposition rates for various particle sizes of aquatic vascular plant detritus have been shown for Phragmites communis Trinius leaves (Hargrave 1972), Spartina alterniflora (Odum and de la Cruz 1967, Gosse-link and Kirby 1974) and Thalassia testudinum (Fenchel 1970). That a similar response was shown for Nereocystis luetkeana and Iridaea cordata detritus indi cates that decomposition rate may be influenced by the amount of surface area exposed to microbial attack. Part of the difficulty in determining a relationship between par ticle size and the parameters tested in Experiments 1 and 2 may be explained by there being only a small amount of crude fibre present in seaweed biomass. Re fractory material accounts for a large proportion of vascular plant detritus, and is composed largely of lignins, celluloses and hemicelluloses which slowly decompose'over a period of months (Lousier and Parkinson 1975, Stachurski and Zimka 1975, 1976 a&b, Gasith and Lawacz 1976). Lignin is the most resistant of these constituents, having a half-life of about one year at 30 C and under optimal conditions for microbial decomposition (Acharya 1935). The amount of lignin present in leaves was given a range of 16-42% by Jensen (1974), who sum marized the work of several authors. With the amount of material lost by lea ching ranging from ca 27-40% (Otsuki and Wetzel 1974, Suberkropp et al. 1976), lignin may account for up to 70% of the particulate fraction of the detritus. In contrast to vascular plants, macrophytic algae contain no lig nin, although their cell walls do contain celluloses and hemicelluloses (Steward 1974) which are moderately resistant. The crude fibre content for the 10 species assayed in this study ranged from 1.2-17.7% of the particulate component. As vascular plants contain a much greater amount of crude fibre than seaweeds it is reasonable to conclude that the rapid decomposition rates of seaweed litter and detritus is at least partially due to a paucity of resistant material. Figures 12a (crude fibre) and 12b (glucose) demonstrate the relationship between decomposition rate and resistant material content; however this parameter was shown to account for only 42.8% of the variance associated with Figure 12a and 39.9% of the variance associated with Figure 12b. Other factors must be involved as well. No doubt some of the variance is due to limitations in the data, but factors determining the resistance and susceptibil ity of seaweeds to attack by microbes are likely to play important roles. The three major structural components of seaweeds have been shown to influence the decomposition process independently. In Experiment 1 soluble matter was isolated from the remaining two components as being preferentially metabolized. From the results of Experiment 2 the crude fibre component was identified as an influence on the decomposition rates of the particulate frac tion; the greater the quantity of crude fibre, the slower the decomposition rate. The three components can thus be ranked in order of soluble, moderately resistant and crude fibre with respect to the ease with which each is metaboliz ed, as has been previously documented for vascular plant material. Detritus Assessment: The accuracy of detritus biomass estimations is questionable. A major criticism is the assumption that the flat, horizontal areas chosen as collecting surfaces are equally as receptive to detritus settlement as uneven surfaces. Such surfaces would be expected to retain a greater proportion of the detritus than the level surfaces. A second criticism is that the biomass data do not take into consideration the decomposition rate of natural detritus. Data in this study indicate that turnover of seaweed detritus is rapid. For Iridaea cordata detritus ca 18-21 days are required, other species requiring a longer turnover time. With sampling intervals of about three weeks, the quan tity of detritus deposited on the bottom in Site 1 will potentially be under estimated by 50%, if its biogenic origin is seaweed biomass. As microscopic - 92 -examination of the detritus determined it to be composed of ca 10% diatomaceous material, the most significant component of phytoplankton in the Strait of Georgia (Hutchinson and Lucas 19 31) , seaweed remains the only source abundant enough to account for the remaining 90% of detritus biomass. It is likely that detritus deposition within Site 1 will be underestimated, perhaps by more than 50%; however, this will not preclude the possibility of making judgements regard ing the fate of detritus formed from seaweed biomass. Faunal Assessment: Vascular plant detritus is a confirmed source of food for fish and invertebrates (Kaushik and Hynes 1968, Iverson 1973, Tenore 1975, Kostalos and Seymour 19 76, Sibert et al. 19 77). Seaweed detritus derived from the phaeo-phyte Dictyopteris zonarioid.es Farlow has been shown to be ingested by the epi-benthic deposit-feeding holothurian Parastichopus parvimensis Clark (Yingst 19 76). Fucus vesiculosus detritus has been utilized by the brine shrimp Acartia tonsa Dana (Roman 1977) and the molluscs Hydrobia ulvae Pennant and Macoma balthica L. (Newell 1975). In each of these experiments seaweed detritus was the only food source such that no indication of the animals' preference for this food re source was attained. In this study Lacuna marmorata, Metacaprella anomala and Cancer oregonensis were delimited as possible respondents to the availability of natural seaweed detritus as a food resource on the basis of the occurrence of at least 75% of their numbers and biomass during the three midsummer faunal collec tions. A critical consideration of these species reveals that Cancer oregonensis is an unlikely respondent as its habit is carnivorous and, furthermore, only 19 individuals were collected such that its qualification may be an artifact of in adequate sampling. Cancer oregonensis will not be considered any further. Inadequate sampling may also be argued as the reason for Meta caprella anomala qualifying as only nine individuals were collected. Although not abundant during the summer of 1976, Metacaprella anomala has been abundant - 93 -at Site 1 in previous years (Foreman unpub.). Circumstantial evidence that Metacaprella anomala is a detritus utilizer was obtained in this study when they were observed attached to the experimental litter bags in numbers from 10-100 individuals about the end of July. This was the only time during the summer of 1976 when their presence was obvious. Lacuna marmorata was very abundant at Site 1 during the summer of 19 76 with about 7400 individuals being collected at densities approaching 70,000/m2 (Figure 16). Of particular significance is the more than 10-fold in crease in the number and dry weight of juvenile Lacuna marmorata individuals during midsummer. Preliminary consideration of other species which may have quali fied at a lesser percentage indicated they were less likely to be significantly dependent on a summer pulse of detritus. Additional species which would have qualified at a 50% acceptance level are Odostomia spp. , Lirularia lirulata Carpenter and Granulina margaritula Carpenter. Odostomia can be removed from consideration as they are generally ectoparasitic in habit (Fretter and Graham 1949). Granulina margaritula and Lirularia lirulata were not particularly abun dant in the faunal collections, did not display strong midsummer peaks in numbers or biomass, presented no evidence of the occurrence of juveniles and as their diets are undocumented it is not possible to discuss their occurrence in relation to the availability of detritus from a positive perspective. An indication that the occurrence of specific marine fauna might be a response to the availability of seaweed detritus as a food source, which was shown in this study to occur about the middle of August, was obtained in the sum mer of 1975 when an extensive 'bloom' of caprellid amphipods, mainly Caprella alaskana Mayer was observed in Site 1. They were estimated to be present at a density of hundreds per square metre. Also evident at this time was a 'scum' of detritus over the bottom, particularly in the central area of the kelp bed. That - 94 -this is at least a periodic phenomenon was confirmed by reference to Foreman's (unpub.) faunal data which contained a record of Caprella alaskana at a density of 520/m2 and Metacaprella anomala at 312/m2 in 1973 and 1972, respectively, and at lesser densities in other years (Table 12). Caine (1977) presents evidence that these two species may utilize detritus. Although neither one is represented in his study, he demon strated (with reference to other authors) that detritus was fed upon by 15 of the 16 species which he investigated. Their mode of feeding was variable, in volving various combinations of filter feeding, scavenging and scraping. Food acquisition was determined to be related to the presence or absence of plumose setae on their second antennae, those species with such antennae obtaining a significant amount of their food by scraping and/or filtering particulate matter. The second antennae of both Metacaprella anomala and Caprella alaskana are characterized by the presence of plumose setae, and since Caine observed that 75% of the stomach contents of caprellids with such setae consisted of diatoms and detritus it is reasonable to conclude that detritus contributes significantly to the diet of these two species. Any argument that Caprella alas kana and Metacaprella anomala are responding to the availability of diatoms is weak. Such a response would have been expected to occur earlier in the year at the time of the spring bloom of diatoms and other phytoplankton in the Strait of Georgia (Hutchinson et al.1929, Gran and Thompson 1930), not during the summer when nutrient levels in the Strait of Georgia are low (Tully and Dodimead 1957). Diatoms comprised only ca 10% of the biomass of detritus samples collected from the substratum in Site 1 during the summer of 1975. No 'bloom' of caprellids was detected within Site 1 during the summer of 1976 when this study was conducted. The scarcity of both Caprella alaskana and Metacaprella anomala during the summer of 1976 cannnot be explained with certainty but it is possible that unusual environmental conditions during Table 12. History of the occurrence (per m ) of two species of Caprel-lidae, Caprella alaskana and Metacaprella anomala, within the summer faunal collections of Dr. R. E. Foreman (unpublished). Foreman's transect units are used, but they are essentially equivalent to the transect units in this study. Number Year Distance along transect jm) C. alaskana M. anomala August 1972 July 1973 July 1975 August 19 75 75 80 60 65 70 75 80 85 95 55 60 85 90 95 50-90 200 2 76 324 456 520 8 4 8 312 28 4 16 8 4 16 several hundred/m * * visual estimation of abundance - 96 -August, generally the warmest month, were at least partially responsible. The Vancouver Weather Office, ca 35 km from Bath Island at Vancouver International Airport, reported August 1976 to be one of the coldest on record. The mean air temperature for August 1976 was 15.9 C. The normal mean air temperature for August is 17.1 C. On only two occasions since 1937 was a lower mean air tempera ture recorded during August. Water temperature was similarly influenced. Based on daily readings near Site 1 (Foreman unpub.) the mean water temperature for the first half of August 1976 was 3.64 C below the mean temperature for August 1975 (12.36 C and 16.2 C, respectively) when Caprella alaskana was very abundant. As metabolism is a function of temperature, persistent cool tem peratures could appreciably lower the growth potential of an organism. Micro bial decomposition rates of seaweed litter would be similarly affected, reducing the quantity of detritus available. Although based only on a visual interpreta tion, the quantity of detritus which accumulated on the bottom during August 1976 was observed to be much less than the quantity observed in 19 75 when a large bloom of Caprella alaskana was observed. In conclusion, it is reasonable to sug gest that the effect of low temperatures on the growth rates of both Caprella alaskana and Metacaprella anomala, coupled with low detritus availability as a food resource, may have been sufficient to prevent a proliferation of these species in 1976. It has not been shown experimentally that Lacuna marmorata util izes detritus as a food source, however E. Cabot (pers. comm.) examined the gut contents of individuals collected near Site 1 and found an abundance of diatoms and amorphous material whose biogenic origin could not be identified. He clas sified this latter material as detritus while conceding it may have been living material rendered unrecognizable due to mastication during and following inges tion. Lacuna are known grazers of seaweeds. Powell (1964) demonstrated that Lacuna fed upon Constantinea subulifera. The author has observed adult Lacuna marmorata grazing upon Nereocystis luetkeana lamina. - 97 -These reports refer only to adult snails. Juvenile snails com prised the bulk of the individuals of Lacuna marmorata collected within Site 1 during the period of maximum detritus availability. Figure 16 demonstrates that 100% of the total number and dry weight of Lacuna marmorata were collected in the zone 30-50 m along the permanent transect. Not only is this the turf community zone (Lindstrom 1973), which aids in the retention of the detritus and provides a habitat for Lacuna marmorata, it is also where maximum detritus biomass was observed during the summer (see also Figure 13, page 73). Based on this evidence, and the results from the simulation of litter and detritus processing which indi cate seaweed detritus to be suitably nutritious for fauna, it does not seem un reasonable to infer that the success of a midsummer recruitment of juvenile Lacuna marmorata individuals is dependent on the availability of seaweed detritus as a food resource. - -SIMJLATION MODEL OF LITTER AND DETRITUS PROCESSING Introduction: To date decomposition models developed to simulate specific as pects of litter and detritus decomposition have been limited to the terrestrial environment. Boling et al. (1975) were primarily concerned with simulating an aspect of leaf and branch litter decomposition by considering the interaction between fractionation of the material by physical abrasion and microbial condi tioning of the resulting particles. Flanagan and Bunnell (1976) developed a model to deal with the influence of moisture, oxygen, temperature and litter composition on the respiration rates of microbes associated with litter, and another to assess the decomposition rates of terrestrial plants under the influ ence of changing substrate quality. The need for this degree of resolution becomes more apparent as the complexity of the system increases. Moisture content, temperature and oxy gen tension within soil can vary daily and seasonally, greatly influencing the decomposition rates of soil borne litter (Nyhan 1976). This requires that they be incorporated into models simulating terrestrial decomposition processes (Hunt 1977, Reuss and Innis 1977). Decomposition rates are also dependent upon the availability of inorganic nutrients, particularly nitrogen (Kaushik and Hynes 1971, Nichols and Keeney 1973, Howarth and Fisher 1976). In a marine system many of these complications can be avoided. The buffering quality of seawater helps alleviate the potential variability in many parameters. There are seasonal variations in the contents of inorganic nitrogen and oxygen in the Strait of Georgia, but it is unlikely their concen trations drop to a level limiting the decomposition rates of the species studied. Oxygen concentrations in the upper 10 m of the Strait of Georgia are consisten tly near 100% saturation (Tully and Dodimead 1957). During litter collections, pockets of litter were occasionally found containing some seaweeds undergoing - 99 -anaerobic decomposition, however, the quantity was insignificant compared to the amount of litter undergoing aerobic decomposition. As seaweed litter tends to retain nitrogen preferentially during the decomposition process, the avail ability of nitrogen is probably not a factor influencing the decomposition rate of most seaweed litter. Substrate quality, temperature, and moisture content remain the major factors to be considered. The effect of substrate quality is accounted for intrinsically within the derived litter decomposition curves leaving temperature the only effect needing to be incorporated into the model. Moisture is obviously not an influential factor. The numerical objectives of the simulation were: 1) to predict the seasonal formation rates, biomass, and longevity of detritus derived from decaying seaweed litter within Site 1 2) to predict the seasonal release rates and quantity of soluble matter released from sea weed litter at Site 1 3) to estimate the nitrogen contents of the detritus formed and soluble matter released from decomposing seaweed litter. Determination of these parameters facilitated a comparison be tween the biomass of detritus predicted to be available as a food resource with in Site 1 and the biomass of detritus obtained from the sample collections. Additionally, an estimate of the seasonal contribution of detritus and soluble matter derived from seaweed litter to the Strait of Georgia was obtained. Model Development: Initially, a four dimensional matrix representing the pool of sea weed litter, the driving variable in the model, was created to permit litter to be referenced in terms of its biogenic origin, the quadrat within the transect from which it was collected, and the location of the transect. Only Fucus distichus, Iridaea cordata, Nereocystis luetkeana (stipe and lamina sections con sidered individually) and Laminaria (L. saccharina and L. groenlandica combined), the species accounting for more than 97% of the quantity of litter collected, - 100 -were incorporated into the model. Extrapolation of the areal profile for each of these species (Figure 2) was facilitated by prorating the 14 month seasonal collections (Figure 5) according to a tenth degree polynomic curve which approx imates the seasonal trend in litter biomass. This curve is presented in Figure 17 for total litter biomass. The litter decomposition curves for these species are presented in Figure 6 (d,g,h,i,j), page 43. For Fucus distichus, Nereocystis luetkeana (stipe) and Laminaria, which decompose exponentially, 1.0% of original dry weight was considered the termination of the decomposition process. The rates were modified by a temperature dependent adjustment factor which accounts for the effect of seasonal temperature differences on decomposition rates. Monthly mean temperatures are presented in Table 13a, based on regular measurements taken at or near Site 1. Temperatures were converted to a decomposition rate adjustment factor (Table 13b) by the following formula, assuming a of 2.0 approximates the effect of temperature on decomposition rates (Boling et al 1975, Reuss and Innis 1977). where : 13.4 - T F = 2 10 F is the decomposition rate adj ustment factor T is the temperature in C The mean temperature during the period when the litter bag experiments were performed was 13.4 C. The adjustment factor was estimated for each day of the year by fitting the following cyclical curve to the adjustment factors determined from the above formula. The formulae for calculation of the following curve are in Croxton et al.(1967). F = 1. 375 + (0.20187 sin(2Tr/366) + 0.29821 cos (2TT/366) ) x I where: F is the decomposition rate adjustment factor I is the day of the year The model was operated over the time period of 28 February 1976, - 101 -Figure 17. Tenth degree polynomic curve fitted to the seasonal biomass data obtained from litter collections along the 95 m transect location within Site 1 from 20 August 19 75 until 2 October 1976. Biomass is in g ash-free dry weight per The curve model is as follows: PB = Z (piDY1_1) ; i = 1,11 where: PB is the predicted litter biomass DY is the day of the year p. are the coefficients l The coefficients are as follows: 1) 0. 3115825195312500E+01 2) -0. 155 3213977813721E+00 3) 0. 5167517066001892E-02 4) -0. 9613301604986191E-•04 5) 0. 1075271284207702E-05 6) -0. 7727231263743306E-•08 7) 0. 3657364633369298E-•10 8) -0. 1130370536756020E- 12 9) 0. 2186710082213890E- 15 10) -0. 2393779959900677E-•18 11) 0. 1127717301078564E-•21 - 102 -Table 13. Mean monthly temperatures (a) and the corresponding decompo sition rate adjustment factor (b) for the period November 1975 until October 19 76. The temperature data are based on periodic readings near Site 1 (Foreman unpub.). See text for an explanation of the adjustment factor. Month a) Temperature (C) b) Adjustment factor January 5.6 1.717 February 6.1 1.659 March 6.4 1.625 April 7.6 1.49May 8.4 1.414 June 11.8 1.117 July 13.4 1.000 August 12.5 1.064 September 13.6 0.986 October 9.6 1.301 November 7.7 1.485 December 6.3 1.636 - 10 3 -when litter biomass was essentially zero, until 31 December 1976. Since no litter collections were made beyond 2 October 1976, data from the autumn of 19 75 were used for the period of October through December 1976. With daily increments beginning on 28 February 1976 litter was mathematically processed according to the temperature corrected specific submodels. Litter biomass available to be decomposed each day was determined by applying the equation for the curve in Figure 17 to the ratio of specific litter:total litter for the most recent samp ling date. The onset of decomposition was delayed by an estimated senescence delay of six days (temperature adjusted) as explained on page 54. Specific litter in each quadrat was processed independently during the simulation. Starting on 28 February 1976, litter which decomposed on this date was subtracted from the litter biomass at the beginning of the day. This calculation was then performed for every subsequent day required to reduce the litter biomass to zero, assuming no further litter deposition. For each of these subsequent days, the remaining litter biomass was subtracted from litter biomass at the beginning of the day to account for daily biomass loss. Remaining litter will be supplemented with freshly deposited litter and undergo decomposi tion on future days. Following performance of this cycle for each species in every quadrat, the data were summed to yield the total quantity of detritus formed and soluble matter released on 28 February 1976, with partial sums for the immediately subsequent days. This entire procedure was then repeated, with daily increments, for the duration of the simulation. During the simulation all soluble matter was released in advance of the particulate material. Until the remaining litter biomass reached the per centage equal to the particulate material content for that species, all export ation was registered as soluble matter. Further decomposition formed detritus. Concomitant with litter decomposition, the nitrogen content of the detritus formed and the soluble matter released was determined. Unfortun-- 104 -ately, due to the rapid decomposition rates of the species involved in the simu lation, minimal nitrogen data were obtained for these species. The data are in sufficient to support firm conclusions but are' suitable for approximating trends for the purpose of modelling. Curve models for estimating litter nitrogen content were selected from those introduced on page 42, with those yielding the best fit being accepted. They are as follows: Fucus distichus Y = -1.09E-02X + 2.83 (L) Iridaea cordata Y 7.04E-0 3X + 1.24 (L) Nereocystis luetkeana (stipe) Y = -9.33E-03X + 2.44 (L) Nereocystis luetkeana (lamina) Y = -2.92E-03X + 5.30 (L) Laminaria Y = 7.85E-05X2 -- 3.39E-02X + 4.92 (Q) where: X is the percentage of litter remaining in the litter bag Y is the percentage nitrogen content of the material remaining in the litter bag The formula derived for calculating the nitrogen content of de tritus and soluble matter is as follows: N. = QL. (PLU. , - PLU.) (PN. + PN. J ( (PN. ,) (PLU. J/PLU.) l 1 1-1 1 l 1-1 1-1 1-1 1_ 2 * ((PN. .) (PLU. _)/PLU.) l-l l-l l - PN. - PN i-1 where: N is the quantity of nitrogen released as soluble matter or as a component of detritus on day i QL is the quantity of litter available for decomposition on day i PLU is the proportion of older litter yet undecomposed (a function of the litter decomposition submodels, Figure 6) PN is the proportion of nitrogen in the litter (a function of the litter nitrogen content submodels listed above) i is a counter for the day during the simulation Detritus decomposition was simulated by usage of the detritus de composition rates obtained for the initial 0-10 day incubation period in Experi ment 2. For Fucus distichus the decomposition rate for 20-30 days was used. All rates were linear and are as follows: Fucus distichus 0.76% per day Iridaea cordata 5.65% per day Nereocystis luetkeana (stipe) 3.12% per day Nereocystis luetkeana (lamina) 3.48% per day Laminaria 2.93% per day - 105 -As the change in nitrogen content of decomposing detritus was not determined, it was modelled as though it decomposed at the same rate as other detritus components. Soluble matter was not decomposed. To reduce the volume of output produced by the simulation, daily incremental data were summed and averaged over 3-4 week intervals. Greater resolution was superfluous and unmanageable. A flow chart outlining the major operations involved in the performance of the simulation is presented in Figure 18. Results: Operation of the simulation model determined the proportional contributions of Fucus distichus, Iridaea cordata, Nereocystis luetkeana and Laminaria to the litter. Table 14 compares the true proportional contributions of each species to their estimated contributions based on sampled litter biomass alone. As expected, the proportional contribution by Fucus distichus was con siderably lower than indicated by the biomass data, due to its particularly slow decomposition rate relative to the other species. The proportional contributions by all other species increased, most dramatically for Nereocystis luetkeana (lamina). The unreliability of litter biomass as an estimator of the true quan tity of litter which undergoes decomposition is apparent. Figure 19 displays the seasonal profile for the rate of detritus formation and release of soluble matter from decomposing seaweed litter within Site 1. Both are seasonal phenomena with peaks occurring during late summer. 2 Maximum observed rates were ca 0.6 and 0.5 g AFDW/m per day for detritus forma tion and soluble matter release, respectively. In total, ca 56% of decomposing litter forms detritus, the remainder being released as soluble matter. Figure 20 displays the predicted detritus biomass formed from litter deposited along the permanent transect location (95 m) in Site 1. Figure 21 presents a similar picture based on total litter deposition within Site 1. - 106 -Figure 18. Flow chart outlining the major operations involved in the simulation of litter and detritus processing within Site 1. LITTER BIOMASS within Site 1: subscripted to species, quadrat, transect and day of the year; as determined from the equation for Figure 17 prorated by the ratio of specific litter:total litter for the immediately preceding sampling date. LITTER NITROGEN CONTENT SPECIFIC SUBMODELS: introduced on page 104. DETRITAL/SOLUBLE MATTER NITROGEN CONTENT CALCULATION: introduced on page 104; requires input from LITTER BIOMASS, LITTER DECOMPO SITION RATE and NITROGEN CONTENT SPECIFIC SUBMODELS.  LITTER DECOMPOSI SPECIFIC SUBMODE in Figure 6, pag TION RATE LS: introduced e 43. LITTER DECOMPOSI TEMPERATURE ADJL LATION: introduc TION RATE STMENT CALCU-ed on page 100. IF the percentage of specific litter biomass processed is less than the equivalent percentage of soluble matter in the litter, soluble matter and soluble nitrogen are released; alternatively, detritus and detrital nitrogen are formed. PREDICTED TOTAL DETRITUS AND DETRITAL NITROGEN BIOMASS formed within Site 1: subscrip ted to species, quadrat, tran sect and day of the year. DETRITUS AND DETRITAL NITROGEN DECOMPOSITION RATE SPECIFIC SUBMODELS: introduced on page 104. PREDICTED DETRITUS AND DETRITAL NITROGEN BIOMASS ACCUMULATION within Site 1. subscripted to quadrat, transect and day of the year. PREDICTED TOTAL SOLUBLE MATTER AND SOLUBLE NITROGEN released within Site 1: subscripted to day of the year. - 10 7 -Table 14. Comparison of the percentage contributions by the major contributors to the litter pool within Site 1 as determined by: a) litter biomass alone b) application of the decomposition rates of these species to litter biomass data. These percentages were determined on an ash-free dry weight basis. Species Fucus distichus Iridaea cordata Nereocystis luetkeana (stipe) Litter biomass 71.98 15.07 1.95 Nereocystis luetkeana (lamina) 7.36 Litter biomass coupled with decomposition rates 40.84 26.22 3.57 23.72 Laminaria 1.69 3.70 Figure 19. Seasonal profiles for the formation rate of detritus and the release rate of soluble matter from decomposing seaweed litter biomass within Site 1. Rates are in g ash-free dry weight per m per day. - 109 -Figure 20. Detritus biomass predicted for the 95 m transect location within Site 1 based on litter collections from that location2 only. Contour intervals are 2.0 g ash-free dry weight per m - 110 -Figure 21. Detritus biomass predicted for Site 1 based on litter collections from all transect locations within Site 1 Contour intervals are 10.0 g ash-free dry weight per - Ill -Reference to Figure 13 (page 73) highlights an obvious discrep ancy between predicted and observed detritus biomass. Based on sampling data, detritus biomass along the permanent transect location peaked at 1.4 g AFDW/n2 9 whereas the predicted quantity was ca 30 g AFDW/ra if all detritus was deposited on the substrate. If detritus biomass is more accurately predicted by incorpor ating all litter data for Site 1, ca 80 g AFDW/m2 is estimated. Accepting that the data incorporated into the model are reasonably accurate, the implication is that detritus accumulation in Site 1 amounts to only 1-5% of the quantity of detritus formed from seaweed litter within Site 1, the remainder being exported. Alternatively, the difference between predicted and observed detritus biomass is a result of litter deposited within Site 1 undergoing decom position elsewhere, its residence time in Site 1 being very short. Three argu ments discount this hypothesis. Most specific litter was collected near stands of the same species. Very little litter was observed outside the seaweed zone. The simulation demonstrated that litter decomposition rates could account for the disappearance of all but 3% of the litter deposited within Site 1. The mean nitrogen content of detritus at the time of its formation was predicted to be 2.48 - 0.03% of its dry weight over the period of the simula tion. The quantity of nitrogen released with the soluble matter was a lesser amount at 1.36 t 0.0 3%. Discussion: Data obtained in this study indicate that ca 80 g AFDW/fa2 of detritus was formed from seaweed litter during 1976. When soluble matter is added to this figure it is increased to 145 g AFDW/n2. With carbon accounting for 50-2 60% of the elemental composition of the organic matter (Round 1965), 70-85 g C/m is the estimate for the amount of carbon leaving seaweed biomass via litter decomposition in Site 1. - 112 -This amount accounts for ca 45% of the quantity of seaweed bio mass lost from the same area as determined from seasonal differences in standing crop biomass (Foreman unpub.). The remaining biomass must be accounted for by detritus formation directly via lamina tip erosion and by Nereocystis luetkeana leaving Site 1 when detached by winds and waves. Johnston et al. (1977) esti mated Laninaria saccharina to lose 40-50% of its gross primary production by lamina tip erosion, a certain percentage of which would be expected to form detritus without being shunted through the litter pool. It must also be considered that seasonal changes in standing crop biomass may inadequately estimate the total quantity of detritus formed from seaweeds. Mann (1972b) estimates the ratio of yearly productionrinitial biomass for populations of Laninaria digitata and Lan inaria longicruris to be 9.8 and 7.2, respectively. Agarum was less productive at 4.2. Thus, without necessarily constituting a major portion of the standing crop biomass within the seaweed zone, these kelps can account for a large portion of the net production. Such an extensive turnover of biomass results in standing crop biomass under estimating total production and subsequent detritus formation and soluble matter release. As Laminaria and Agarum are characteristic of both Site 1 and Mann's (1972b) system this consideration is probably appropriate; however, the indications are that Nereocystis luetkeana has the highest biomass turnover of the plants within Site 1 (Foreman unpub.). As Nereocystis luetkeana was rank ed tenth in 'importance' of the 'significant' species within Site 1 and third in its contribution to the litter pool, changes in biomass may not severely underestimate total detritus formation and soluble matter release when the entire system is considered. This is supported by Foreman's seasonal biomass data which indicate that 45% of the biomass loss occurred in the depth range 0-3 m below mean sea level. This is the zone dominated by Fucus distichus and Iridaea - 113 -cordata, the two dominant contributors to the litter pool. A large biomass turnover has not been shown to be characteristic of these species. The re maining biomass loss is accounted for by the other eight 'significant' species most of which are found in the depth range of 10-30 m below mean sea level. Of these, only Nereocystis luetkeana and possibly Laminaria are characterized by high biomass turnover. The litter and detritus biomass data (Figures 5, and 20 and 21, respectively) indicate the peak period of detritus formation from seaweed occurs during late summer. This would be consistent with a hypothesis that maximum productivity occurs during the summer months, based on Mann's (1972a) interpretation of the results of Krey (1967) and Sutcliffe (1972) which imply a peak in particulate material biomass derived from seaweed during early spring, at the time of maximum seaweed productivity in St. Margaret's Bay (Mann 1972a). As only 1-5% of detritus predicted to have been formed from seaweed litter, and a lesser percentage of total detritus formed from seaweed biomass, accumulated within Site 1, the majority of seaweed detritus must be processed elsewhere. Webster et al. (1975) collected an amount of organic matter equivalent to 15% of total plant production in sediment traps placed at deep stations (60 and 65 m ) in St. Margaret's Bay, Nova Scotia. Data from shallower stations were less reliable. During the year of their study Laminaria produc tion alone exceeded phytoplankton production by three fold, and with the major settlement peaks occurring when plankton production was low, they conclude seaweed detritus to be the most likely origin of the organic matter collected. Data from this study indicate that < 5% of seaweed detritus is deposited within the seaweed zone, leaving at least 80% to be exported, and subsequently decom posed in coastal waters, along with the soluble matter released. Lenz (1977) obtained results which may be considered evidence - 114 -of the presence of seaweed detritus in coastal water. In an attempt to show a positive correlation between the standing crop biomasses of phytoplankton and/or zooplankton and that of detritus in the Kiel Bight, West Baltic Sea, only data from stations below 15 m depth supported his hypothesis. In water above 15 m depth negative (although nonsignificant) correlations were obtained. The suggestion is that the detritus is of an allochthonous nature, contrary to Lenz's hypothesis that it was formed autochthonously. Sources such as air borne dust, coastal erosion and sediment were discounted but seaweeds were not referenced. Seaweeds are a normal feature of the Western Baltic coastline, and with the Kiel Bight being an enclosed area a possible explanation of his results has been overlooked. Odum and de la Cruz (196 7) determined a maximum rate of 1.4 g AFDW/m^ per day for the exportation of organic matter from an east coast estu arine salt marsh. The average daily rate of detritus formation from seaweed litter is in the range of 0.2-0.4 g AFDW/m2, but the total amount formed may be at least double this figure when complemented by detritus from erosive pathways. If these data are typical, detritus formed from seaweeds should exceed contri butions from other plant systems unless such systems are more abundant than the seaweed zone. In the Strait of Georgia, where the seaweed zone is a marked feature of the coastline, this is not the case. There are major estuarine salt marshes at the mouths of the Fraser and Squamish Rivers, but they account for a small proportion of the total coastline. Eelgrass (Zostera marina) mea dows are also present in the Strait of Georgia, near Robert's Bank (Forbes 1972, Moody 1978) and Nanaimo (Foreman 1975, Sibert et al. 1977). Rates of formation of Zostera marina detritus are not available for the ecosystem level but there is no evidence to suggest they will be significantly higher than those obtained for the salt marsh systems. It is unlikely that detritus originating from either - 115 -system will exceed the quantity originating from seaweed biomass other than in the immediate vicinity of the respective systems. The ecological roles of seaweed detritus and vascular plant detritus will be dissimilar due to the composition of the biomass undergoing decomposition. Seaweed detritus appears to be too short-lived and only sea sonally available to provide a long term food resource for fauna. Alterna tively, vascular plant detritus has been documented as a long term food resource for fauna during periods when primary production is low (Darnell 1967b). The predicted nitrogen content of seaweed detritus, determined in this study to be ca 2.48% of its dry weight, is probably underestimated. This is partially due to the specific submodels for the species incorporated in to the simulation (page 104) generating less rapid increases in the relative nitrogen content of decomposing litter than indicated by the trend in Figure 7 (page 58). Additionally, the simulation decomposed detrital nitrogen at the same rate as other detrital components. This is probably an underestimation of its true decomposition rate when considering the pattern observed for litter decomposition. To obtain an indication of the suitability of seaweed detritus as a food resource for fauna a C:N ratio was estimated for the detritus formed. Generic and/or class estimates of the elemental carbon contents, as a percen tage of dry weight, for the five species modelled are as follows: Nereocystis luetkeana ca 20% (J. Whyte pers. comm.) Fucus spiralis and Fucus vesiculosus 33-36% (Vinogradov 1953, Niell 1976) Laminaria 12-27% (Vinogradov 1953, Niell 1976) Rhodophyta (in general) 20-38% (Niell 1976) These data were prorated according to the percentage contribution by each species to detrital biomass to yield a C:N ratio of 10-13:1. This is less than the value of 17:1 which Russell-Hunter (1970) considers the minimum nitrogen content rendering a food resource suitably nutritious for most fauna. The C:N - 116 -ratio of the soluble matter released is in the range of 18-24:1 and must be considered nutritively poor. As the C:N ratio for detritus is probably an overestimation, it follows that the ratio for soluble matter is an under estimation. In comparison, vascular plant detritus usually undergoes a considerable degree of processing before it attains a nutritive value that renders it suitable for consumption by potential consumers. Harrison and Mann (19 75b) found that between 35 and 102 days were required for microbes to re duce the C:N ratio of Zostera marina detritus from an initial value of 20.2:1 to less than 17:1. Iverson (1973) performed preference experiments which dem onstrated that decomposing leaves were not fed upon until nitrogen enrichment occurred. In this study Lacuna marmorata, Caprella alaskana and Meta-caprella anomala were delimited as possible utilizers of natural seaweed detritus, based on their morphology, habit, spatial and/or temporal distribu tion patterns. The implication was that these species may be responding to a summer pulse in the availability of sufficiently nutritious seaweed detritus. It is necessary that experiments be performed to determine these species' food preference, and their growth and survival while utilizing this resource, in order to conclude with certainty that they can respond to the availability of seaweed detritus as a food resource. - \n-SUMMATION Previous examinations of the role of organic detritus in coastal ecosystems have consistently underplayed the significance of the con tribution by detritus originating from seaweed biomass (Darnell 1967b, Fenchel 1972, 1973, Perkins 1974). That detritus derived from seaweed biomass may contribute significantly to coastal energetics was first seriously considered by Mann (1972a). This study supports the interpretation that seaweed detritus biomass exported to coastal waters is likely the major macrophytic source of particulate organic material for the Strait of Georgia, British Columbia, and perhaps exceeds the contribution from planktoriic sources during non-bloom periods. It is reasonable to extrapolate that the particulate material con tent of enclosed areas characterized by a seaweed zone (e.g. St. Margaret's Bay, Kiel Bight) receives a significant contribution from seaweed biomass. The annual quantity of seaweed detritus formed and soluble matter released from the system studied is estimated to be at least 45% derived from seaweed litter, with a maximum rate of detritus formation being observed during late summer. This amount is complemented by detritus formed directly via erosion of kelp lamina tips. Decomposition experiments indicated that seaweed litter decom poses very rapidly, seaweed detritus is short-lived, and that this was at least partially due to its paucity of structural material resistant to metabol ism by microbes. This has probably been a reason for underestimations of seaweed detritus and soluble matter contributions to total coastal organic material relative to other coastal macrophytes based on sampled biomass alone. The inability to distinguish adequately between organic material originating from phytoplankton and seaweed biomass further complicates this problem (Sutcliffe 1972, Webster et al. 1975) . - 118 -Previous authors' unawareness of the degree to which seaweed detritus biomass contributes to coastal food resources has precluded an interpretation of the importance of this resource to benthic and pelagic faunal distribution patterns. This study has confirmed that seaweed detritus is suitably nutritious for fauna, having a C:N ratio of 10-13:1 or less. Sea weed detritus is thus more acceptable than living seaweed biomass which has C:N ratios ranging from 13.8:1 to 27.2:1 for Laminaria (Mann 1972a) and 40:1 to 80:1 for kelps in general (Russell-Hunter 1970). Although this study could not conclude with certainty that seaweed detritus is a food resource relied upon by specific benthic fauna, there is circumstantial evidence that some species are at least periodically dependent upon its availability. The implication is that the quantity and relatively high nitrogen content of seaweed detritus renders it particularly suitable as a food resource such that one must expect that it has a very significant role in the structure and function of coastal ecosystems. - uq-LITERATURE CITED Acharya, C.N., 19 35: Studies on the anaerobic decomposition of plant materials. JJJ. Comparison of the course of decomposition of rice straw under anaerobic, aerobic and partially aerobic conditions. Biochem. 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Lee, 1977: Effect of meiofauna on incorpora tion of aged eelgrass, Zostera marina, detritus by the polychaete Nephthys incisa. J. Fish. Res. Board Can. 34:563-567 - 126 -Tully, J.P., and A.J. Dodimead, 1957: Properties of the water in the Strait of Georgia, British Columbia and influencing factors. J. Fish. Res. Board Can. 14:241-319 Vinogradov, A.P., 1953: The elementary chemical composition of marine organ isms. Mem. Sears Found. Mar. Res. No. 2, 647 pp Webster, T.J.M., M.A. Paranjape and K.H. Mann, 1975: Sedimentation of organic matter in St. Margaret's Bay, Nova Scotia. J. Fish. Res. Board Can. 32:1399-1407 Whyte, J.N.C., and J.R. Englar, 1975: Basic organic chemical parameters of the marine alga Nereocystis luetkeana over the growing season. Fish. Res. Board Can. Tech. Rep. 589, 42 pp Widdowson, T.B., 1973: The marine algae of British Columbia and northern Washington: revised list and keys. Part I. Phaeophytes (brown algae). Syesis 6:81-96 Widdowson, T.B., 1974: The marine algae of British Columbia and northern Washington: revised list and keys. Part II. Rhodophyceae (red algae). Syesis 7:143-186 Wolff, T., 1976: Utilization of seagrass in the deep sea. Aqua. Bot. 2:161-174 Yingst, J.Y., 1976: The utilization of organic matter in shallow marine sedi ments by an epibenthic deposit-feeding holothurian. J. Exp. Mar. Biol. Ecol. 23:55-69 Zobell, C.E., 1971: Drift seaweeds on San Diego County beaches. In: W.J. North (Ed.), The biology of giant kelp beds (Macrocystis) in California: 269-314. Nova Hedwigia 32 (Suppl.) - 12 7 -APPENDIX I A) Numerical species code for litter assessment data in Appendix I (B,C,D). B) Litter assessment data for seasonal collections at 95 m within Site 1. C) Litter assessment data for collections at 5, 35, 65 and 95 m within Site 1 on either 27 July or 3 August 1976. D) litter assessment data for the collection at Site 2 on 10 November 1975. A. 01 Plocamium coccineum var. pacificum 02 Gigartina papillata 03 Fucus distichus 04 ' Rhodomela larix 05 Odonthalia floccosa 06 Iridaea cordata 07 Nereocystis luetkeana (stipe) 08 Nereocystis luetkeana (lamina) 09 Laminaria saccharina 10 Laminaria groenlandica 11 Constantinea subulifera 12 Ulva spp./Monostroma spp. 13 Prionitis lanceolata Harvey 14 Sargassum muticum (Yendo) Fensholt 15 Agarum spp. 16 Zostera marina 17 Costaria costata (Turner) Saunders 18 Laurencia spectabilis Postels and Ruprecht 19 Laminaria spp. 20 Rhodymenia palmata (L.) Greville 21 Halymenia spp. 22 Analipus japonicus (Harvey) Wynne 2 3 Gracilariopsis sjoestedtii (Kylin) Dawson 24 Enteromorpha spp. 25 Ceramium spp. 26 Cryptopleura ruprechtiana (J. Agardh) Kylin 27 Gelidium spp. 28 Gigartina spp. 29 Microcladia borealis Ruprecht 30 Rhodymenia pertusa (Postels and Ruprecht) J. Agardh 31 Gymnogrongus linearis (Turner) J. Agardh 32 Alaria spp. 33 Porphyra torta Krishnamurthy 34 Gloiosiphonia capillaris (Hudson) Carmichael 35 Fauchea lanciniata J. Agardh 36 Rhodoptilum plumosum (Harvey and Bailey) Kylin 37 Bossiella spp. 38 Pterosiphonia bipinnata (Postels and Ruprecht) Falkenberg 39 Desmarestia viridis (Muller) Lamouroux 40 Polyneura latissima (Harvey) Kylin 41 Callophyllis flabellulata Harvey 42 Bonnemaisonia nootkana (Esper) Silva 43 Gigartina exasperata Harvey and Bailey 99 Unidentified litter - 128 -WET DRY ASH-FREE LOCATION SPECIES WEIGHT WEIGHT DRY WEIGHT DATE QUADRAT (G/10W2) (G/10M2) (G/10M2) 1 20/08/75 ' 1 95 10 -20 03 3.5400 0. 6950 0.4650 2 20/08/75 ' 1 95 20 -30 03 24.6050 3.9450 2.8650 3 20/08/75 ' 1 95 20 -30 14 0. 4400 0.0859 0.0457 4 20/08/75 J 95 20 -30 18 1.0050 0. 1616 0.1052 5 20/08/75 ' 1 95 20 -30 12 0.1150 0. 017 1 0.0107 6 20/08/75 ' 1 95 20 -30 13 0.2200 0. 069 1 0.0407 7 20/08/75 ' 1 95 30 -40 07 0.3550 0.0367 0.0252 8 20/08/75 1 95 30 -40 14 1.7700 0.2740 0. 1593 9 20/08/75 " 1 95 40 -50 07 133.9301 19. 6850 10.6700 10 20/08/75 1 1 95 40 -50 08 275.9500 28. 1050 16.2350 11 20/08/75 ' 1 95 40 -50 06 404.7849 84.2100 51.2200 12 20/08/75 1 95 40 -50 19 110.9500 13.0650 8.4050 13 20/08/75 ' 1 95 40 -50 15 18.0500 2.7100 1.5350 14 20/08/75 ' 1 95 40 -50 05 8.0500 1.9150 1.0850 15 20/08/75 ' 1 95 40 -50 32 3.4400 0. 3592 0.2023 16 20/08/75 ' 1 95 40 -50 12 10.8050 1.9350 1. 2750 17 20/08/75 ' I 95 40 -50 18 6.6650 0. 8702 0.4911 16 20/08/75 1 95 40 -50 14 0.8100 0. 1258 0.0713 19 20/08/75 ' 1 95 40 -50 26 0.5200 0.0098 0.0 044 20 2Q/08/75 ' 1 95 40 -50 44 0.2400 0. 0256 0.0155 21 20/03/75 " 95 50 -60 03 316.0300 39.9350 24.0850 22 20/08/75 ' 1 95 50 -60 07 489.5400 60. 5550 36.9200 23 20/03/75 • 1 95 50 -60 19 98.3850 18.5900 9.1000 24 20/03/75 ' 1 95 50 -60 06 7.9600 0.9600 0. 5501 25 20/08/75 ' 95 50 -60 13 0.0800 0. 0097 0. C049 26 20/03/75 ' 95 50 -60 18 0.8200 0.0094 0.0060 27 20/C6/75 ' 1 95 50 -60 15 1.4550 0. 2059 0.1107 26 20/08/75 ' 1 95 50 -60 17 2.1000 0. 258 1 0.1491 29 20/08/75 " 95 50 -60 21 0. 7700 0. 0958 0.0583 3C 20/08/75 ' 1 95 60 -70 07 332.5100 43.2400 24.5650 31 20/08/75 ' 1 95 60 -70 0 3 342.2148 42.5750 25.8650 32 20/08/75 ' 1 95 60 -70 19 64 .9350 11.3650 6.4200 33 20/08/75 " 95 60 -70 21 2.1650 0. 2593 0.1608 34 20/08/75 " 95 60 -70 32 2.0250 0.3252 0.1973 35 20/08/75 ' 1 95 60 -70 12 7.6100 1.8600 1.0850 36 20/08/75 ' 1 95 70 -80 07 62.1850 6.3430 3.C050 37 20/0 8/75 ' 95 70 -80 08 187.8149 21.0950 13.0900 38 20/08/75 • 95 70 -80 19 68.1050 8. 3400 5. 3 6 00 39 20/08/75 ' 1 95 70 -80 06 0.1250 0. 0148 0.0091 40 20/08/75 ' 1 95 30 -90 07 132.0900 13. 6600 9.0600 41 20/08/75 ' 1 95 80 -90 08 125.2050 12.9400 8.0700 42 20/0 8/75 ' 1 95 80 -90 14 0.7400 0.1592 0.0982 43 20/03/75 ' 1 95 80 -90 19 0.3900 0. 0446 0.0236 44 20/08/75 ' 1 95 90- 100 07 85.6050 12. 0700 7.0050 45 20/08/75 ' 1 95 90- 100 08 133.4850 14.3700 8.815C 46 20/08/75 ' 1 95 90- 100 12 5.5100 0.7350 0.4850 47 20/08/75 ' 1 95 90- 100 19 14.4050 1.9150 1.0850 48 02/09/75 ' ! 95 00 -10 07 10.6000 0.9300 0.4620 49 02/09/75 ' 1 95 20 -30 03 191.1400 52.9150 39.9000 50 02/09/75 ' 1 95 20 -30 06 6.3200 1.0400 0.6912 51 02/09/75 ' 1 95 20 -30 12 1. 9700 0. 3138 0.2020 52 02/09/75 ' I 95 20 -30 07 0.5800 0.0669 0.0464 53 02/09/75 ' 1 95 20 -30 13 2. 1500 0.4837 0.3166 54 02/09/75 1 1 95 20 -30 04 0.7390 0. 1219 0.0788 55 02/09/75 ' 1 95 20 -30 28 0.3467 0. 1017 0.0517 -129-56 02/09/75 1 95 20 -30 14 0.1428 0. 0450 0.0319 57 02/09/75 1 95 20 -30 18 0.0893 0.0225 0.0 147 58 02/09/75 1 95 20 -30 01 0.1867 0. 0435 0.0311 59 02/09/75 1 95 30 -40 03 86.4250 22.9030 16.4900 60 02/09/75 1 95 30 -40 06 6.5150 1.5850 1. 0 900 61 02/09/75 1 95 30 -40 14 3. 2900 0.5950 0. 4225 62 02/09/75 1 95 30 -40 13 0.9300 0. 1849 0.1176 63 02/09/75 1 95 30 -40 18 7.3050 0.8750 0. 6550 6a 02/09/75 1 95 30 -40 12 6.2350 1.0300 0.6226 65 02/09/75 1 95 30 -40 07 0.2585 0.0350 0.0212 66 02/09/75 1 95 30 -40 08 0.2145 0.0350 0.0199 67 02/09/75 1 95 30 -40 05 0.1892 0. 0470 0. 0291 68 02/09/75 1 95 40 -50 19 163.8250 28. 6450 13.9150 69 02/09/75 1 95 40 -50 06 177.6851 38.5400 27.0650 70 02/09/75 1 95 40 -50 07 406.4399 1 4. 4750 8.1700 71 02/09/75 1 95 40 -50 08 124.0600 3.0250 1.8300 72 02/09/75 1 95 40 -50 17 8.8750 1.0100 0.4850 73 02/09/75 1 95 40 -50 18 1.7550 0.2300 0.1521 74 02/09/75 1 95 40 -50 16 2.4650 0. 2900 0. 1872 75 02/09/75 1 95 40 -50 12 32.4100 5.3050 3. 5800 76 02/09/75 1 95 40 -50 18 7. 8900 1. 3700 0.9050 77 02/09/75 1 95 5 0 -60 07 5.5250 0.7726 0.4500 78 02/09/75 1 95 50 -60 08 6.2000 0. 8397 0.5051 79 02/09/75 1 95 50 -60 28 0.6 950 0.2063 0. 1006 80 02/09/75 1 95 50 -60 12 7.0000 1 .3150 0.8450 81 02/09/75 1 95 50 -60 19 7. 8300 5.0300 2.5700 82 02/09/75 1 95 50 -60 06 85.3900 12.7150 9.0950 83 02/09/75 1 95 60 -70 15 49.0550 7.3200 4. 0650 84 02/09/75 1 95 60 -70 06 17.9800 3.6400 2.5450 85 02/09/75 1 95 6 0 -70 07 99.1050 8.4450 5. 3200 86 02/09/75 1 95 60 -70 08 26.9900 3. 1100 1.7700 87 02/09/75 1 95 6 0 -70 15 8.6150 1. 4850 0.8300 88 02/09/75 1 95 60 -70 17 2.2300 0.2250 0.1162 89 02/09/75 1 95 60 -70 18 0.7417 0.0430 0.2930 90 02/09/75 1 95 60 -70 01 0.0437 0.0071 0.C049 91 02/09/75 1 95 60 -70 12 0.4406 0.0705 0.0459 92 02/09/75 1 95 60 -70 20 1.1362 0. 1256 0.0656 93 02/09/75 1 95 70 -80 19 19.9100 2.8450 1.4050 94 02/09/75 1 95 70 -80 03 10.0000 2.3850 1.8150 95 02/09/75 1 95 70 -80 07 67.2600 7.3500 4. 3550 96 02/09/75 1 95 70 -80 08 45.6300 4. 8150 2.8700 97 02/09/75 1 95 70 -80 12 12.0400 1.5150 0.9750 98 02/09/75 1 95 70 -80 21 8.8900 1. 3450 0.9350 99 02/09/75 1 95 70 -80 16 0.2935 0. 0271 0.0170 1 00 02/09/75 1 95 80 -90 14 0.3370 0. 0412 0.0281 101 02/09/75 1 95 80 -90 07 71.4450 8.0600 4.2500 102 02/09/75 1 95 80 -90 08 71.0200 6.8750 3.9550 103 02/09/75 1 95 80 -90 03 6.2100 1. 3950 1.0350 1 04 02/09/75 1 95 80 -90 12 9.1550 1.3300 0.8700 105 02/09/75 1 95 80 -90 14 1.3217 0.2309 0.1592 106 02/09/75 1 95 80 -90 19 23.7550 2.6500 1.3400 107 02/09/75 1 95 80 -90 21 1 .8252 0.2312 0.1610 108 02/09/75 1 95 80 -90 18 0.3439 0.0392 0.0270 109 02/09/75 1 95 90- 100 08 43.2150 3.7600 2.1100 1 10 02/09/75 1 95 90- 100 12 4.1100 0.8450 0.5150 1 11 02/09/75 1 95 90- 100 06 0.0538 0.0040 0.0023 112 02/09/75 1 95 90- 100 16 0.7178 0.0626 0.0400 1 13 02/09/75 1 95 90- 100 23 0.2474 0.0531 0.0322 1 14 04/10/75 1 95 10 -20 07 12.3900 1.4100 0.8300 1 15 04/10/75 1 95 20 -30 03 6.1 100 1.3400 1.0200 - 130 -116 04/10/75 ' 1 95 20 -30 07 0.8700 0. 0950 0. 0518 1 17 04/10/75 ' 1 95 20 -30 12 0.1150 0.0181 0. 0T19 1 18 04/10/75 1 95 40 -50 07 60.9600 7.2450 4. 3 250 119 04/10/75 1 1 95 40 -50 08 83.1600 9. 6650 5. 6300 120 04/10/75 ' 1 95 40 -50 06 19.2900 5. 1450 3. 7 150 121 04/10/75 ' 1 95 40 -50 12 10.1500 1. 8900 1. 2000 122 04/10/75 ' 1 95 40 -50 18 9.4900 1. 3700 0. 9450 123 04/10/75 * 95 50 -60 07 262.1899 42.4400 23. 5800 124 04/10/75 1 1 95 50 -60 08 30.9300 3.8550 2. 2550 125 04/10/75 " 1 95 50 -60 19 12.9100 2. 6950 1. 3600 126 04/10/75 * 1 95 50 -60 15 6.5450 1. 1450 0. 6250 127 04/10/75 ' 1 95 50 -60 06 13.9450 3.4800 2. 4750 128 04/10/75 " 1 95 50 -60 12 3.6850 0.5350 0. 3291 129 04/10/75 ' 1 95 60 -70 07 1 14.9350 17.7100 10.2800 130 04/10/75 1 95 60 -70 08 72.3450 9.2500 5. 5300 131 04/10/75 ' 1 95 60 -70 19 1.2200 0. 1800 0. 1024 132 04/10/75 1 95 60 -70 12 2.3250 0. 3800 0. 2303 133 04/10/75 " I 95 60 -70 16 0.2228 0. 021 0 0. 0127 134 04/10/75 1 95 70 -80 08 38.9950 4.3850. 2. 6 150 135 04/10/75 ' 1 95 70 -80 07 100.8450 13.0300 7. 4350 136 04/10/75 " 1 95 70 -80 12 5.9800 1.0500 0. 6585 137 0 4/10/75 ' 1 95 70 -80 21 1.4557 0.1303 0. 0910 138 04/10/75 ? 95 70 -80 19 19.4400 2. 6600 1. 3750 139 04/10/75 ' 1 95 80 -90 19 108.1300 20. 2000 10. 3550 140 04/10/75 ' 1 95 80 -90 07 11.2000 1.3600 0. 8350 141 04/10/75 1 95 80 -90 08 23.5900 3.2100 1. 8650 142 04/10/75 1 95 90-100 07 55.7900 5.1950 2. 7350 143 04/10/75 ' 1 95 90-100 08 33.8800 3.6400 2. 2 6 50 144 04/10/75 ' 1 95 90-100 16 0.3900 0. 0635 0. 0409 145 04/10/75 ' 1 95 90-100 03 0.7050 0. 1465 0. 1 069 146 04/10/75 ' 1 95 90-100 14 0.4450 0. 1070 0. 0755 147 04/10/75 ' 1 95 90-100 12 0.8600 0. 1765 0. 1 170 148 04/10/75 ' 1 95 90-100 19 3.1100 0.4755 0. 2560 149 04/10/75 1 95 90-100 25 0.2549 0. 0289 0. 0129 150 04/10/75 1 95 90-100 24 0.0909 0. 007 3 0. 0 035 151 09/11/75 ! 95 10 -20 03 5.0100 0. 8650 0. 6600 152 09/11/75 ' 1 95 30 -40 07 34.8600 4.7850 3. 4300 153 09/11/75 1 95 30 -40 16 0.5560 0. 0577 0. 0351 154 09/11/75 1 95 30 -40 18 0.8350 0. 1046 0.0685 155 09/11/75 ' 1 95 40 -50 06 0.4977 0. 0980 0. 0682 156 09/11/75 ' 1 95 40 -50 08 4.9605 0.4310 0. 2520 157 09/11/75 ' 1 95 40 -50 07 225.6000 32.0000 17. 4800 158 09/11/75 1 95 40 -50 12 0.5860 0. 0893 0. 0585 159 09/11/75 1 95 40 -50 19 1.8 114 0.2257 0. 1201 160 09/11/75 ! 95 50 -60 07 106.3600 15. 8550 9. 0 100 161 09/11/75 1 95 50 -60 05 1.0163 0. 2070 0. 1 281 162 09/11/75 1 95 50 -60 06 1.2730 0.3269 0. 2370 163 09/11/75 1 95 60 -70 07 652.5750 99.5700 48. 5550 164 09/11/75 • 1 95 60 -70 08 11.9618 1. 1097 0. 6444 165 09/11/75 1 95 60 -70 15 8.0589 3. 2286 1. 8150 166 09/11/75 1 95 60 -70 12 0.5 117 0.0870 0. 0606 167 09/11/75 1 95 60 -70 19 1.8080 0.2693 0. 1 378 168 09/11/75 1 95 70 -80 08 17.0100 1.8850 1. 1000 169 09/11/75 ' 1 95 70 -80 07 39.0100 6. 0805 3. 3594 170 09/11/75 1 95 70 -80 19 1.9064 0.2548 0. 1280 171 09/11/75 ' 1 95 70 -80 12 0.5899 0.0915 0. 0613 172 09/11/75 1 95 80 -90 08 15.4150 1.3650 0. 7821 173 09/11/75 ' 1 95 80 -90 07 78.8750 7.3100 4. 4250 174 09/11/75 1 95 80 -90 • 19 13.6350 1.7300 0. 8524 175 09/11/75 1 95 90-100 07 73.4850 7. 5550 4. 0200 -. 131 176 09/11/75 1 95 90- 100 08 177 09/11/75 1 95 90- 100 06 178 09/11/75 1 95 90- 100 18 179 09/11/75 1 95 90- 100 26 180 09/11/75 ' I 95 90- 100 16 181 10/12/75 ' I 95 30 -40 08 182 10/12/75 ' I 95 30 -40 18 183 10/12/75 ' I 95 30 -40 16 184 10/12/75 1 I 95 30 -40 03 185 10/12/75 ' 95 40 -50 07 186 10/12/75 * 95 40 -50 08 187 10/12/75 1 95 40 -50 12 188 10/12/75 ' I 95 40 -50 03 189 10/12/75 ' I 95 40 -50 13 190 10/12/75 ' I 95 50 -60 07 191 10/12/75 ' I 95 50 -60 18 192 10/12/75 ' I 95 50 -60 03 193 10/12/75 ' I 95 50 -60 12 194 10/12/75 ' I 95 50 -60 08 195 10/12/75 ' 95 60 -70 07 196 10/12/75 ' I 95 60 -70 16 197 10/12/75 ' I 95 60 -70 19 198 10/12/75 ' I 95 60 -70 08 199 10/12/75 H 95 70 -80 08 200 10/12/75 ' I 95 70 -80 07 201 10/12/75 I 95 70 -80 19 202 10/12/75 ' I 95 70 -80 06 203 10/12/75 1 95 7 0 -80 12 204 10/12/75 ' I 95 70 -80 13 205 10/12/75 1 95 80 -90 07 206 10/12/75 ' 1 95 80 -90 08 207 10/12/75 ' 1 95 80 -90 12 208 10/12/75 ' 1 95 80 -90 19 209 10/12/75 ' 1 95 90- 100 08 210 10/12/75 ' 1 95 .90- 100 07 211 10/12/75 ' 1 95 90- 100 12 212 16/01/76 ' 1 95 30 -40 07 213 16/01/76 ' 1 95 50 -60 07 214 16/01/76 ' 1 95 50 -60 16 215 16/01/76 ' 1 95 60 -70 07 216 16/01/76 ' 1 95 70 -80 07 217 16/01/76 ' 1 95 80 -90 07 218 16/01/76 ' 1 95 90- 100 07 219 28/02/76 ' 1 95 30 -40 32 220 28/02/76 1 95 50 -60 32 221 28/02/76 1 95 80 -90 07 222 14/03/76 ' 1 95 20 -30 03 223 14/03/76 ' 1 95 20 -30 12 224 14/03/76 1 95 40 -50 16 225 14/03/76 1 95 40 -50 18 226 14/03/76 1 95 40 -50 26 227 14/03/76 ' 1 95 40 -50 12 228 14/03/76 ' 1 95 40 -50 33 229 14/03/76 1 95 40 -50 19 230 14/03/76 ' 1 95 40 -50 03 231 14/03/76 ' 1 95 40 -50 31 232 14/03/76 ' 1 95 50 -60 19 233 14/03/76 1 95 50 -60 30 234 14/03/76 ' 1 95 50 -60 12 235 14/03/76 \ 95 50 -60 18 12.6350 1. 0600 2. 0 150 0.9593 0. 2815 0. 2023 0.4932 0. 075 2 0. 0549 0.9806 0. 2638 0. 1701 0.5208 0. 1235 o. 0784 2.2775 0. 2320 0.1359 0.2334 0. 026 5 0. 0179 0.9128 0. 0884 0. 0556 1.0582 0. 2279 0. 1731 119.4450 6. 4950 3. 1250 4.6847 0. 3865 0. 2253 0.6439 0. 097 1 0. 0518 4. 1573 1. 1567 0. 7032 0.0621 0. 006 2 0. 0042 31.4500 2. 8400 1. 8400 0.1078 0. 0135 0. 0100 2.8950 0.7572 0. 5 572 0.0571 0. 0090 0.0590 7.6800 0. 7192 0. 4325 73.3900 6. 6200 4. 6200 0.9771 0. 1003 0. 0641 1.0150 0. 1 175 0. 0612 1.9200 0. 1798 0. 1 039 25.7900 2. 6700 1. 5700 65.5350 4. 3500 2. 6650 3.8498 0. 4575 0. 2369 0. 9337 0. 1858 0. 1 405 0. 4 138 0. 0617 0. 0373 0.0911 0.0288 0.0155 57.4550 7. 2650 4. 0 150 12.7201 1. 3770 0. 7991 1.9476 0. 4317 0. 2925 0. 2939 0. 0455 0. 0235 7.1852 0. 727 1 0. 4500 63.8800 3. 6650 1. 9550 0.4747 0. 0855 0. 0549 8.2700 1. 9150 1. 2750 15.5100 2. 6900 1. 4250 0.5458 0. 0633 0. 3993 51.9050 6. 4650 3. 5 100 34.8800 5. 2800 3. 4100 79.0600 11. 7350 9. 2850 67.2750 9. 1300 4. 5800 22.5050 4. 0750 2. 4500 12.4850 2. 2650 1. 4150 13.7050 2. 1550 1. 3350 17.8300 3. 2100 2. 3900 0.7910 0. 1609 0. 1086 0.2738 0. 0473 0. 0316 0.9937 0. 1304 0. 0914 0.5304 0. 0924 0. 0622 4.3806 0. 2620 0. 1772 3.7850 • 0. 8650 0. 3490 5.1550 0. 6750 0. 3840 4.4800 0. 8600 0. 6249 0.1194 0. 0229 0. 0 137 13.5950 1. 9850 1. 0200 0.1392 0. 0298 0. 0 157 2.7204 0. 4863 0. 3219 0.7566 0. 1380 0. 0960 - 132 -236 14/0 3/76 1 1 95 50 -60 33 1.5219 0. 3209 0.1363 237 14/03/76 1 1 95 60 -70 19 33.3650 3. 4900 1.7900 238 14/03/76 ' 1 95 60 -70 12 4.4150 0.6050 0.4278 239 14/03/76 ' 1 95 60 -70 26 0.7586 0.1240 0.0822 240 14/03/76 ' 1 95 60 -70 23 0.7683 0.0613 0.0400 241 14/03/76 " 95 70 -80 19 3. 5262 0. 8484 0.4551 242 18/04/76 " 1 95 1 0 -20 06 33.5350 6.3150 4. 6000 243 18/04/76 ' 1 95 10 -20 03 3.9550 0.9200 0.7182 244 18/04/76 " 1 95 10 -20 12 1.3579 0. 1828 0. 1251 245 18/04/76 ' 1 95 10 -20 01 0.1 107 0.0227 0.0 151 246 18/04/76 ' 1 95 10 -20 34 0.4318 0. 0667 0.0435 247 18/04/76 ' 1 95 20 -30 06 0. 1964 0. 0448 0.0306 248 18/04/76 ' 1 95 30 -40 12 0.9776 0. 1829 0.1385 249 18/04/76 ' 1 95 30 -40 18 0.0486 0. 0113 0.0078 250 18/0 4/76 ' 1 95 30 -40 35 0.4103 0.0799 0.0545 251 18/04/76 ' 1 95 30 -40 36 0.0078 0.0050 0.0038 252 18/04/76 ' 1 95 40 -50 37 0.6554 0.5 100 0.0720 253 18/0 4/7 6 1 95 40 -50 38 0.2803 0. 042 1 0.0254 254 18/04/76 1 95 40 -50 12 0.3239 0.0622 0.4720 255 18/04/76 ' 1 95 40 -50 19 2.2882 0.431 5 0.2497 256 18/04/76 • 1 95 50 -60 07 2.8460 0. 2307 0.0894 2 57 18/0 4/76 ' 1 95 50 -60 08 3.2728 0.3249 0. 1882 258 18/0 4/76 1 1 95 50 -60 12 3.7472 0.5568 0.3665 259 18/04/76 ' 1 95 50 -60 16 0.8896 0. 1290 0.0825 260 18/04/76 ' 1 95 50 -60 01 0.3636 0. 0524 0.0371 261 18/04/76 ' 1 95 50 -60 06 0.0931 0. 0146 0.0102 262 18/04/76 ' 1 95 60 -70 08 0.0709 0. 0142 0.0 102 263 18/04/76 ' 1 95 60 -70 12 0.5282 0. 0845 0.0610 264 18/04/76 1 95 60 -70 99 0.0509 0. 0 168 0.0073 265 18/04/76 ' 1 95 70 -80 08 0.2627 0. 0420 0.0252 266 18/04/76 1 95 70 -80 07 0.2610 0.0250 0. 0109 267 18/04/76 ' 1 95 70 -80 16 0.3000 0.0347 0.0221 268 18/04/76 • J 95 80 -90 12 2.0972 0. 3388 0. 2303 269 18/04/76 ' 95 80 -90 19 1.9176 0.4783 0.2449 270 18/04/76 • 1 95 90- 100 08 0.9496 0.0968 0.0558 271 18/04/76 " 1 95 90- 100 03 7.5500 1. 3400 0.9851 272 18/04/76 ' 1 95 90- 100 19 6.7250 0.7950 0.5001 273 13/05/76 ' 1 95 20 -30 03 33.1000 5.5850 4.0319 274 13/05/76 ' 1 95 20 -30 12 2.7590 0.3973 0.2761 275 13/05/76 ' 1 95 30 -40 12 1.0404 0. 2515 0.1991 276 13/05/76 J 95 30 -40 01 3.9977 0. 7109 0.4999 277 13/05/76 ' 1 95 40 -50 19 9.0550 1.2250 0.7850 278 13/0 5/76 ' 1 95 40 -50 05 4.6226 0. 6578 0.4434 279 13/05/76 1 95 40 -50 06 0.6 114 0. 1356 0.0974 280 13/05/76 " 1 95 40 -50 30 0.8033 0. 1418 0.0924 281 13/05/76 ' 1 95 40 -50 39 0. 4857 0.0716 0.0488 282 13/05/76 " 1 95 40 -50 40 0.4606 0.0773 0.0522 283 13/05/76 ' 1 95 40 -50 12 2.4051 0.3856 0.2720 284 13/05/76 1 95 40 -50 01 0.3726 0.0574 0.0360 285 13/05/76 ' 1 95 40 -50 26 0.3832 0. 083 1 0.0581 286 13/05/76 " 1 95 40 -50 08 0. 8949 0.1187 0.0768 287 13/05/76 1 95 40 -50 03 6.2472 1. 4914 1.1313 288 13/05/76 1 95 50 -60 08 15.8850 1.7550 0.8450 289 13/05/76 1 95 50 -60 12 1.9601 0.2736 0.1696 290 13/0 5/76 ' 1 95 50 -60 06 1.5374 0.3080 0.2148 291 13/05/76 " 1 95 50 -60 01 0.9067 0.1276 0.0756 292 13/05/76 1 95 50 -60 17 4.6083 0. 4984 0.2685 293 13/05/76 ' 1 95 50 -60 18 0.7720 0. 1075 0.0620 294 13/05/76 " 1 95 50 -60 32 2.0154 0.4179 0.1991 295 13/05/76 1 95 50 -60 36 0.3038 0. 0562 0.0242 - 133 -296 13/05/76 " I 95 6 0-70 12 3.8245 0.4063 0.2288 297 13/05/76 " I 95 60- 70 19 6.3100 0. 721 1 0.4 162 298 13/0 5/76 1 95 60- 70 08 6.3 182 0. 6107 0.2747 299 13/05/76 1 95 70- 80 19 60.1800 6.8250 3. 9050 300 13/05/76 ' 1 95 70- 80 08 14.5000 1.6200 0.8800 301 13/05/76 ' 1 95 70- 80 21 0.9292 0. 2889 0.2023 302 13/05/76 " 1 95 70- 80 12 8.3423 0.9848 0.5673 303 13/05/76 1 95 80- 90 32 4.8430 0.7709 0.5019 304 13/05/76 ' 1 95 80- 90 08 2.2702 0.2221 0.1154 305 13/05/76 < 1 95 8 0-90 07 0.7471 0. 0873 0.0455 306 13/05/76 • 1 95 80- 90 17 0.9479 0.0558 0.0300 307 13/05/76 1 95 90-100 08 5.9302 0. 5884 0.2707 308 27/05/76 ' 1 95 20- 30 03 23.2200 3.7100 2.2250 309 27/0 5/76 • 1 95 3 0-40 08 16.6071 3.0046 1.0604 310 27/05/76 " 1 95 30- 40 06 1.1753 0. 2220 0.1148 311 27/05/76 ' 1 95 30- 40 12 0.9594 0. 2078 0.1218 312 27/05/76 ' 1 95 40- 50 12 2.0550 0.2802 0.1687 313 27/05/76 ' I 95 40- 50 07 0.7146 0. 0560 0.0258 314 27/05/76 ' 1 95 40- 50 08 166.0100 19.3800 9. 1350 315 27/05/76 1 95 40- 50 21 0.2598 0. 0269 0.0142 316 27/05/76 " 1 95 40- 50 39 2.2138 0. 2408 0. 1261 317 27/05/76 " 1 95 40- 50 05 3.0646 0.5944 0.3627 318 27/05/76 " 1 95 40- 50 06 7.2250 1. 5250 0.6750 319 27/05/76 1 95 40- 50 30 2.6233 0.4636 0. 2867 320 27/05/76 ' 1 95 40- 50 18 4.1712 0.6603 0.3398 321 27/05/76 ' 95 40- 50 99 8.8300 1. 6100 0. 7250 322 27/05/76 • 1 95 40- 50 28 2. 2480 0. 1582 0.0773 323 27/05/76 " 1 95 40- 50 19 97.6350 13.7150 7.7 100 324 27/05/76 ' 1 95 50- 60 08 394.7700 41.5150 20.1500 325 27/05/76 ' 1 95 50- 60 03 53.4600 11.2350 8.3050 326 27/05/76 ' 1 95 50- 60 17 26.5650 2. 9600 1.7 2 00 327 27/05/76 1 95 50- 60 26 2.6600 0.5650 0. 4100 328 27/05/76 " 1 95 50- 60 12 18.5000 2.2850 1.5350 329 27/05/76 ' 1 95 50- 60 19 162.4150 19.9450 12.6600 330 27/0 5/76 ' 1 95 50- 60 06 32.9950 6. 1750 4. 2350 331 27/05/76 1 95 50- 60 13 8.3400 1.2650 0. 8400 332 27/05/76 " 1 95 50- 60 21 4.3400 0. 8400 0.6500 333 27/05/76 ' 1 95 5 0-60 05 14.2650 2. 6250 1. 4150 334 27/05/76 1 95 50- 60 28 4.8000 1.0700 0.8050 335 27/05/76 " 1 95 50- 60 18 13.2500 1.9300 1.0800 336 27/05/76 ' 1 95 50- 60 41 1.5500 0.2700 0. 1750 337 27/05/76 " 1 95 50- 60 99 1 5. 5350 2.9000 1.3300 338 27/0 5/7 6 1 95 60- 70 19 6.3724 0. 7580 0.4949 339 27/05/76 1 95 60- 70 12 1 .3167 0. 162 1 0.0983 340 27/05/76 ' 1 95 60- 70 05 0.0697 0.0120 0.0080 341 27/05/76 1 95 6 0- 70 08 109.5050 10. 1850 4.9050 342 27/05/76 " 1 95 60- 70 07 0.2621 0.0214 0.0111 343 27/05/76 • 1 95 70- 80 07 40.3400 5.3600 3.5550 344 27/05/76 • 1 95 70- 80 08 56.5000 5.735 0 3.1100 345 27/05/76 1 95 70- 80 12 5.1486 0.6758 0.4 133 346 27/05/76 I 95 70- 80 39 1. 4528 0. 1386 0.0738 347 27/05/76 1 95 70- 80 16 0.6 961 0.0794 0.0521 348 27/05/76 ' ! 95 80- 90 08 61.9600 5.815 0 3. 0450 349 27/05/76 ' 1 95 80- 90 07 3.0702 0. 4064 0.2312 350 27/05/76 1 95 80- 90 12 50.0400 5. 1600 3.4700 351 27/05/76 1 95 80- 90 19 3.6000 0. 5200 0. 3500 352 27/05/76 J 95 50- 60 17 1.7334 0.1906 0.1218 353 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2300 3.16 50 392 16/06/76 1 95 60 -70 07 11. 0400 1.5600 0.9700 393 16/06/76 1 95 60 -70 12 0. 9285 0.1 136 0.0677 394 16/06/76 1 95 60 -70 06 2. 3621 0. 4602 0.2811 395 16/06/76 1 95 70 -80 08 52. 6150 6. 0200 5. 1000 396 16/06/76 1 95 70 -80 07 14.4000 2. 1950 1.4750 397 16/06/76 1 95 70 -80 19 2. 9389 0. 3173 0. 1807 398 16/06/76 1 95 80 -90 07 78. 7900 10.6300 6.5700 399 16/06/76 1 95 80 -90 08 26. 8700 2. 9200 1.8650 400 16/06/76 1 95 80 -90 12 0. 2207 0. 0640 0.0423 401 16/06/76 1 95 80 -90 26 1. 5169 0.2424 0.1331 402 16/06/76 1 95 80 -90 21 0. 6624 0.0974 0.0629 403 . 16/06/76 1 95 8 0 -90 32 1 . 5529 0.2156 0.1379 404 16/06/76 1 95 80 -90 39 2. 9886 0.4057 0.2240 405 16/06/76 1 95 90- 100 07 20. 5400 2.0950 1.3450 406 16/06/76 1 95 90- 100 08 24.9850 2.3850 1.3650 407 16/06/76 1 95 90- 100 03 4. 8500 0.9800 0.4450 408 07/07/76 1 95 20 -30 19 4. 3446 0. 7538 0.5292 409 07/07/76 1 95 20 -30 08 0. 3828 0.051 1 0.0335 410 07/07/76 1 95 30 -40 06 10. 5881 1.7868 1.2479 411 07/07/76 1 95 30 -40 26 1. 9508 0.281 3 0.2076 412 07/07/76 1 95 30 -40 12 0. 2 424 0.0299 0.0215 413 07/07/76 1 95 40 -50 08 313. 4749 36.9900 23.4950 4 14 07/07/76 1 95 40 -50 06 54. 9250 8. 9000 6.0650 415 07/07/76 1 95 40 -50 19 3. 8012 0. 536 1 0.3453 -. 13b -416 07/07/76 1 95 40 -50 18 1.2201 0. 1341 0.0806 417 07/07/76 1 95 4 0 -50 12 7.1478 0.9584 0.7075 4 18 07/07/76 1 95 50 -60 0.8 224.8300 27.1650 18.2450 419 07/07/76 1 95 5 0 -60 19 94.0550 11.5650 5.9450 420 07/07/76 1 95 60 -70 12 86.6050 10.9550 6.9950 421 07/07/76 1 95 60-70 08 130.9 150 13.4500 8.5750 422 07/07/76 1 95 60 -70 07 71.7900 4. 1100 2.2350 423 07/07/76 1 95 6 0 -70 19 153.0950 17.9750 10.9100 424 07/07/76 1 95 70 -80 07 114.4100 8.7150 5. 1250 425 07/07/76 1 95 70 -80 19 56.6400 6.4400 3.4350 426 07/07/76 1 95 70 -80 18 0.3661 0.0470 0.0273 427 07/07/76 1 95 70 -80 08 253.8101 26.3700 16.6950 428 07/07/76 1 95 80 -90 19 28.7000 1.8350 1.2050 429 07/07/76 1 95 8 0 -90 17 13.1600 2.9150 1.6350 430 07/07/76 1 95 80 -90 06 0.8025 0.1617 0.1129 431 07/0 7/76 1 95 80 -90 07 20.3550 2. 6550 1. 6500 432 07/07/76 1 95 80 -90 08 114.0950 12. 1100 7.6300 433 07/07/76 1 95 90- 100 19 1.0981 0. 1423 0.0700 434 07/07/76 1 95 90- 100 07 3.4135 0. 341 5 0.2058 435 07/07/76 1 95 90- 100 08 91.0650 9.4400 5.7500 436 27/07/76 1 95 20 -30 03 5.5710 1.1911 0.8128 437 27/0 7/76 1 95 20 -30 19 5. 1550 0.6734 0.4718 438 27/07/76 1 95 20 -30 12 2.8377 0.3288 0.1991 439 27/07/76 1 95 20 -30 08 6.4184 0. 6369 0. 4476 440 27/07/76 1 95 20 -30 18 5. 1469 0.5896 0.3410 441 27/07/76 1 95 3 0 -40 03 101.5150 18.8700 15.2200 442 27/07/76 1 95 30 -40 06 v7.7150 1. 3300 0.9150 443 27/07/76 1 95 30 -40 18 12.1050 1. 1250 0. 7850 444 27/07/76 1 95 30 -40 08 25. 7850 2.1600 1.4800 445 27/07/76 1 95 30 -40 19 8.2500 1.3100 1.0850 446 27/07/76 1 95 30 -40 12 16.7500 1.9050 1.5950 447 27/0 7/76 1 95 40 -50 08 570.7849 47. 2100 33. 8850 448 27/07/76 1 95 40 -50 07 26.5200 2.0750 1.5150 449 27/07/76 1 95 40 -50 19 25.8300 3. 1900 1.8350 450 27/07/76 1 95 40 -50 06 37.8 100 5.3700 4.0650 451 27/07/76 1 95 40 -50 01 2.6062 0. 3195 0.1724 452 27/07/76 1 95 40 -50 18 1.5821 0. 1304 0.0890 453 27/07/76 1 95 40 -50 21 2.1989 0. 2472 0.1563 454 27/07/76 1 95 40 -50 12 11. 4400 1.2000 1.0050 455 27/07/76 1 95 40 -50 03 7.6900 1. 2900 1.0150 456 27/07/76 1 95 50 -60 08 108.6100 13.7900 9.1550 457 27/07/76 1 95 50 -60 07 31.0800 5.5550 3.6750 458 27/07/76 1 95 50 -60 19 52.5500 18. 7700 15.6050 459 27/07/76 1 95 50 -60 12 8.5800 1.5450 1.2200 460 27/07/76 1 95 50 -60 21 9.0000 1.1600 0.9600 461 27/07/76 1 95 50 -60 16 0.9015 0. 1039 0.0568 462 27/07/76 1 95 60 -70 08 205.9950 21. 1850 13.9850 463 27/07/76 1 95 6 0 -70 07 71.4000 10.3550 7.2100 464 27/0 7/76 1 95 60 -70 12 23. 3565 0. 3875 0.2426 465 27/0 7/76 1 95 60 -70 19 9.5694 1.0723 0.5183 466 27/07/76 1 95 70 -80 08 8.4700 0.9550 0.7700 467 27/07/76 1 95 70 -80 07 34.9300 5.2000 3. 4250 468 27/07/76 1 95 70 -80 32 14.1950 2. 1800 1.5650 469 27/07/76 1 95 8 0 -90 03 5.2741 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7850 507 18/08/76 ' 1 95 50 -60 07 1.3701 0. 1348 0. 0914 508 18/08/76 ' 1 95 60 -70 08 191.6700 21.4300 14. 6600 509 18/08/76 ' 1 95 60 -70 07 17.0700 2.7150 1 . 7600 510 18/08/76 1 95 60 -70 12 2.3436 0. 271 1 0. 1 897 511 18/08/76 ' 1 95 60 -70 17 14. 2900 5.7050 2. 3450 512 18/08/76 ' 95 70 -80 07 34.6350 5.0950 3. 2250 513 18/08/76 ' 1 95 70 -80 08 35.1 150 4. 1850 2. 1 400 514 18/08/76 1 95 70 -80 12 0.7280 0. 1040 0. 0832 515 18/0 8/76 ' 1 95 70 -80 19 2.0366 0. 276 1 0. 1289 516 18/08/76 ' 1 95 70 -80 18 0.1193 0. 0152 0. 0097 517 18/08/76 " 1 95 80 -90 07 64.4550 10.0350 6. 5900 518 18/08/76 1 95 80 -90 08 2.4 27.8 0. 3023 0. 1847 519 18/08/76 • 1 95 80 -90 19 5.9485 0.7400 0. 4008 520 18/08/76 ' 1 95 90- 100 07 23.0500 3.6700 2. 5250 521 18/0 8/7 6 1 95 90- 100 08 6.3524 0. 7209 0. 4139 522 18/08/76 ' 1 95 90- 100 21 0.346 1 0.0468 0. 0307 523 18/08/76 ' 1 95 90- 100 17 16.9200 2.8200 1. 5300 524 12/09/76 1 95 20 -30 08 23.2000 2.4700 1. 4400 525 12/09/76 1 95 20 -30 14 10.6000 1.8350 0. 9850 526 12/09/76 1 95 30 -40 14 69.6250 10.0350 5. 7150 527 12/09/76 ' I 95 30 -40 03 94.9050 19.6250 14.1200 528 12/09/76 • 1 95 30 -40 06 93.5450 19.3950 13. 2500 529 12/09/76 1 95 30 -40 19 106.8550 19.7150 14. 6200 530 12/09/76 1 95 30 -40 08 242.0400 27.2200 16. 9900 531 12/09/76 1 95 30 -40 12 49.2850 7.6000 5. 0550 532 12/09/76 1 95 30 -40 4 1 21.3200 2.0650 1. 7150 533 12/09/76 1 95 30 -40 18 37.2750 5.8450 2. 7900 534 12/09/76 1 95 30 -40 26 11.6500 2.2050 1. 1950 535 12/09/76 1 95 40 -50 08 494.0750 53.0150 33. 1 850 - 137 -536 12/09/76 1 95 40 -50 07 6.7943 0.7635 0. 4728 537 12/09/76 1 95 40 -50 12 110.7150 16. 3100 11. 2500 538 12/09/76 1 95 40 -50 26 16.8000 2.8600 1. 7800 539 12/0 9/76 1 95 40 -50 19 424.7200 70. 0050 49. 4300 540 12/09/76 1 95 40 -50 03 167.2500 30. 9400 22. 5900 541 12/0 9/76 1 95 40 -50 18 35.0950 4. 8500 2. 5200 542 12/09/76 1 95 40 -50 43 26.4500 4.0750 2. 7300 543 12/09/76 1 95 40' -50 06 94.0550 18.1000 12. 4500 544 12/09/76 1 95 40 -50 21 45.6350 6.4750 4. 3200 545 12/09/76 1 95 40' -50 14 2.5160 0. 3758 0. 2594 546 12/09/76 1 95 40 -50 01 3.0232 0.3604 0. 1587 547 12/09/76 1 95 40 -50 41 7.0569 0.8647 0. 4507 548 12/09/76 1 95 50 -60 19 163.5000 23.5150 15.1800 549 12/09/76 1 95 50 -60 08 100.5550 10.8800 6. 7650 550 12/09/76 1 95 50 -60 12 24.4950 4.0650 2. 6050 551 12/09/76 1 95 50 -60 06 12.2150 2.7050 1. 8050 552 12/09/76 1 95 50 -60 05 0.9159 0.2738 0.1134 553 12/09/76 1 95 50 -60 13 0.2111 0.0845 0. 0 4 82 554 12/09/76 1 95 50 -60 26 7.1013 1.3148 0. 8489 555 12/09/76 1 95 50 -60 07 7.0305 0. 8064 0. 4694 556 12/09/76 1 95 50 -60 18 5.1796 0. 8388 0. 3636 557 12/09/76 1 95 50 -60 41 8.9100 1. 2250 0. 4850 558 12/09/76 1 95 50 -60 01 4. 6600 0. 6750 0. 4600 559 12/09/76 1 95 5 0 -60 03 5.7300 1.3850 0.3000 560 12/0 9/76 1 95 60 -70 08 94.4600 10. 7150 6. 6150 561 12/09/76 1 95 6 0 -70 07 1.8890 0. 2702 0. 1686 562 12/09/76 1 95 60 -70 19 50.1850 7. 2950 4. 8100 563 12/09/76 1 95 60--70 12 14.5500 2.0600 1. 2700 564 12/09/76 1 95 70 -80 07 49.1700 9. 1950 5. 8850 565 12/09/76 1 95 70 -80 08 70.5900 8. 1500 5. 2500 566 12/09/76 1 95 70 -80 32 3.1765 0.5835 0. 3575 567 12/09/76 1 95 70 -80 12 3.4576 0.4796 0. 3367 568 12/09/76 1 95 70 -80 01 0.4577 0. 0929 0. 0407 569 12/09/76 1 95 70 -80 18 0.6422 0. 1059 0. 0579 570 12/09/76 1 95 70 -80 03 1.2381 0. 2448 0. 1720 571 12/09/76 1 95 70 -80 16 1.2396 0. 1758 0. 1249 572 12/09/76 1 95 70 -80 19 0.3588 0. 0595 0.0338 573 12/09/76 1 95 80 -90 08 29.1750 2. 9950 1. 7300 574 12/09/76 1 95 80 -90 12 0. 7766 0. 1930 0. 1 180 575 12/09/76 1 95 80 -90 07 29.2950 4.2350 2. 5950 576 12/09/76 1 95 80 -90 16 0.4 123 0.0803 0. 0479 577 12/09/76 1 95 80 -90 13 0.6259 0. 1630 0. 0806 578 12/09/76 1 95 80 -90 17 1.7170 0.2763 0. 1578 579 12/09/76 1 95 80 -90 19 2.8581 0.4127 0. 2221 580 . 12/09/76 1 95 90- 100 17 5.0999 0. 641 3 0. 3992 581 12/09/76 1 95 90- 100 08 3 .9500 0.4527 0. 2720 582 12/09/76 1 95 90- 100 07 6.5977 1. 0538 0. 7129 583 12/09/76 1 95 90- 100 12 0.0684 0.0172 0. 0 126 584 02/10/76 1 95 20 -30 03 77.0000 17.8100 12. 1050 585 02/10/76 1 95 20 -30 12 2.9621 0.6880 0. 5202 586 02/10/76 1 95 20 -30 06 8.2050 2.5350 1. 8100 587 02/10/76 1 95 20 -30 14 3.6172 0. 8046 0. 4312 588 02/10/76 1 95 20 -30 18 1.4642 0. 3212 0. 1 420 589 02/10/76 1 95 20 -30 05 0.2628 0. 1382 0. 0954 590 02/10/76 1 95 30 -40 19 250.6851 47.4450 33. 9450 591 02/10/76 1 95 30 -40 08 547.3850 52.4300 31. 7250 592 02/10/76 1 95 30 -40 06 133.7350 35.0600 24. 8400 593 02/10/76 1 95 30 -40 26 36.4550 7. 8300 4. 5650 594 02/10/76 1 95 30 -40 28 12.1550 2.5100 1. 3950 595 02/10/76 1 95 30 -40 03 48.5350 10.4250 7. 5450 - 138 -596 02/10/76 1 95 30 -40 05 11.8950 3. 8100 2. 4800 597 02/10/76 1 95 30 -40 18 51.2 05 0 8. 0500 3. 7950 598 02/10/76 1 95 30 -40 12 56.9350 8. 3750 5. 6 400 599 02/10/76 1 95 40 -50 19 271.4250 53. 1700 39. 3750 600 02/10/76 1 95 40 -50 08 356.4199 32. 8100 19. 4250 601 02/10/76 1 95 40 -50 12 32.4350 5. 1750 3. 5450 6 02 02/10/76 1 95 40 -50 03 15.6200 4. 4700 2. 9450 603 02/10/76 1 95 40 -50 14 10.4550 1. 8350 1. 0950 604 02/10/76 1 95 40 -50 26 13.4 150 2. 9800 1 . 8500 605 02/10/76 1 95 40 -50 18 1.9460 0. 3532 0. 17 75 606 02/10/76 1 95 40 -50 06 17.5400 4. 6700 3. 0650 607 02/10/76 1 95 50 -60 08 248.4351 22.0700 12. 5000 608 02/10/76 1 95 50 -60 07 64.6100 4. 9000 2. 5300 609 02/10/76 1 95 50 -60 12 0.6199 0. 1287 0. 0917 6 10 02/10/76 1 95 50 -60 03 3.6947 0. 7197 0. 5092 6 11 02/10/76 1 95 50 -60 19 34.4100 4. 2600 2. 6 100 612 02/10/76 1 95 60 -70 07 85.7700 7. 7900 4. 2400 6 13 02/10/76 1 95 60 -70 08 29.2000 4. 2600 2. 4050 6 14 02/10/76 1 95 60 -70 03 12.1650 2. 3900 1. 7200 6 15 02/10/76 1 95 6 0 -70 19 2.7 120 0. 2795 0. 1468 6 16 02/10/76 1 95 60 -70 12 2.2073 0. 5147 0. 2864 6 17 02/10/76 1 95 70 -80 07 135.8199 27.4400 14. 8400 618 02/10/76 1 95 70 -80 08 42.5650 4. 4700 2. 4300 619 02/10/76 1 95 70 -80 32 10.5100 0.5750 0. 3700 620 02/10/76 1 95 70 -80 19 7.6 931 0. 9847 0. 4548 621 02/10/76 1 95 80 -90 08 17.4650 1. 9950 0.9450 622 02/10/76 1 95 80 -90 07 43.5850 7. 6800 4. 3200 623 02/10/76 1 95 80 -90 19 7.2600 0. 9400 0. 4650 624 02/10/76 1 95 90- 100 08 6.0332 0. 5654 0. 2909 625 02/10/76 1 95 90- 100 07 26.5850 4. 3300 2. 3450 - 139 -WET DRY ASH-FREE LOCATION SPECIES WEIGHT WEIGHT DRY WEIGHT DATE QUADRAT (G/IOM2) (G/10W2) (G/10/f2) 1 03/08/76 1 05 20 -30 03 484.6650 88.6950 68.0250 2 03/08/76 1 05 20 -30 06 1140.0000 338. 5798 227.5250 3 03/08/76 1 05 20 -30 26 16.4550 2.7000 1.7650 4 03/08/76 1 05 20 -30 08 7.6588 0.8080 0.5588 5 03/08/76 1 05 20 -30 18 12.6600 1.4950 1.0150 6 03/08/76 1 05 20 -30 14 2.7200 0.3694 0.2613 7 03/08/76 1 05 20 -30 28 2.1704 0.5108 0.3060 8 03/0 8/76 1 05 20 -30 05 28.5400 4.9250 3.0600 9 03/08/76 1 05 20--30 19 15.7750 2.4950 1. 9350 10 03/08/76 1 05 20 -30 04 21.5350 3.5000 2.4050 11 03/08/76 1 05 20 -30 12 44.2800 5.7100 3.7150 12 03/08/76 1 05 30 -40 06 7.1800 1.5450 0.8750 13 03/08/76 1 05 30 -40 12 9.2500 1.9250 1.1050 14 03/08/76 1 05 30 -40 13 3.2001 0.5774 0.4277 15 03/08/76 1 05 40 -50 06 320.5850 69.7750 51.6050 16 03/08/76 1 05 40 -50 03 104.6800 20.8350 12.4600 17 03/08/76 1 05 40 -50 26 8.4350 1. 8100 1. 2750 18 03/08/76 1 05 40 -50 28 0.9000 0.2450 0.1750 19 03/08/76 1 05 40 -50 12 13.8050 2. 3150 1.4200 20 03/08/76 1 05 40 -50 05 12.3300 2.2100 1 . 3600 21 03/08/76 1 05 40 -50 01 10.1650 1.4200 0.8800 22 03/08/76 1 05 40 -50 17 2.5400 0.3200 0. 2150 23 0 3/0 8/76 1 05 4 0 -50 04 0.0933 0. 020 1 0.0111 24 03/08/76 1 05 40 -50 18 1. 2958 0. 1641 0.0911 25 03/08/76 1 05 40 -50 13 3.9750 0.9650 0.5950 26 03/08/76 1 05 40 -50 19 1. 9 421 0.3449 0.2367 27 03/08/76 1 05 40 -50 21 0.2569 0. 0330 0.0177 28 03/08/76 1 05 50 -60 06 81.3450 15.0000 11.4100 29 03/08/76 1 05 50 -60 12 16.5950 2.5400 1.3350 30 03/08/76 1 05 50 -60 08 25.5950 2.3450 1.4450 31 03/08/76 1 05 50 -60 07 4.6602 0.7386 0.5394 32 03/08/76 1 05 50 -60 03 12.7750 2.8100 2.1950 33 03/08/76 1 05 50 -60 21 2.5422 0.2946 0.1662 34 03/0 8/76 1 05 50 -60 18 1.3132 0. 1538 0.0790 35 03/08/76 1 05 50 -60 26 0.0822 0.0118 0.0061 36 03/08/76 1 05 50 -60 19 35.7500 0. 0464 0.0284 37 03/08/76 1 05 60 -70 08 35.6100 3.5950 2.2700 38 03/08/76 1 05 60 -70 19 6.2579 0.8419 0.4437 39 03/08/76 1 05 60 -70 14 6.2900 0.9050 0.6400 40 03/08/76 1 05 60 -70 21 1.9572 0.2372 0.1358 41 03/08/76 1 05 60 -70 06 1.1912 0. 188 1 0.1023 42 03/08/76 1 05 70 -80 17 36.7900 4.2850 2.5350 43 03/08/76 1 05 70 -80 08 152.0551 16.5450 10.3250 44 03/08/76 1 05 70 -80 19 73.9400 10.0150 6.3350 45 03/08/76 1 05 70 -80 03 8.8400 1.5800 0.9900 46 03/08/76 1 05 70 -80 04 0.0239 0.0080 0.0050 47 03/0 8/76 1 05 70 -80 12 0.2479 0.0419 0.0246 48 03/08/76 1 05 80 -90 08 85.1200 8. 5000 5.2650 49 03/08/76 1 05 80 -90 06 4.5886 0.8636 0.3296 50 03/08/76 1 05 80 -90 19 75.2100 7.6600 4.6950 51 03/08/76 1 05 80 -90 03 1.2371 0. 248 3 0.1741 52 03/08/76 1 05 90- 100 19 16.5350 2.6100 1.6150 53 03/08/76 1 05 90- 100 06 16.5450 2.4800 1.4950 54 03/08/76 1 05 90- 100 08 188.2300 19.9550 12.1750 55 03/08/76 1 05 90- 100 07 53.6950 6.2000 3.6750 56 03/08/76 ' I 35 20- 30 57 03/08/76 ' I 35 20- 30 58 03/08/76 ' I 35 2 0-30 59 03/08/76 " I 35 20- 30 60 03/08/76 ' I 35 20- 30 61 03/08/76 1 35 20- 30 62 03/08/76 1 35 20- 30 63 03/08/76 ' 1 35 20- 30 64 03/08/76 ' J 35 30- 40 65 03/08/76 1 35 30- 40 66 03/08/76 1 35 30- 40 67 03/08/76 ' 1 35 30- 40 68 03/08/76 1 35 30- 40 69 03/08/76 ' 1 35 30- 40 70 03/08/76 1 35 30- 40 71 03/08/76 ' 1 35 30- 40 72 03/08/76 1 35 30- 40 73 03/08/76 ' I 35 40- 50 74 03/08/76 * 1 35 40- 50 75 03/08/76 • 1 35 40- 50 76 03/08/76 ' 1 35 40- 50 77 03/08/76 ' 1 35 50- 60 78 0 3/0 8/76 ' 1 35 5 0-60 79 03/08/76 ' 1 35 50- 60 80 03/08/76 1 35 60- 70 81 03/08/76 -1 35 60- 70 82 03/08/76 ' 1 35 6 0-70 83 03/08/76 ' 1 35 60- 70 84 03/08/76 1 35 70- 80 85 03/08/76 ' 35 70- 80 86 03/08/76 ' 1 35 80- 90 87 03/08/76 ' 1 35 80- 90 88 03/08/76 1 35 80- 90 89 03/08/76 ' 1 35 80- 90 90 03/08/76 • 1 35 90- 100 91 03/08/76 ' 1 35 90-100 92 03/08/76 1 65 20- 25 93 03/08/76 ' 1 65 20- 25 94 03/08/76 ' 1 65 20- 25 95 03/08/76 " ! 65 20- 25 96 03/08/76 1 65 20- 25 97 03/08/76 1 65 20- 25 98 03/08/76 ' 1 65 30- 40 99 03/08/76 ' 1 65 30- 40 100 03/08/76 1 65 50- 60 101 03/03/76 1 65 50- 60 102 03/08/76 1 65 50- 60 103 03/08/76 1 65 50- 60 104 03/08/76 1 65 60- 70 105 03/08/76 ' 1 65 60- 70 106 03/08/76 1 65 60- 70 107 03/08/76 ' 1 65 70- 80 108 03/08/76 1 65 70- 80 109 03/08/76 1 65 70- 80 1 10 03/08/76 1 65 70- 80 1 1 1 03/08/76 1 65 70- 80 112 03/08/76 1 65 70- 80 1 13 03/08/76 ] 65 70- 80 1 14 03/08/76 1 65 80- 90 1 15 03/08/76 1 65 80- 90 140 -03 373.0649 76.5350 56.0650 08 17. 8650 1. 7500 1.2850 06 49.2500 10.5650 7.6650 05 11.3616 2.002 1 1.1739 12 2.7305 0.4110 0.2495 28 1.8208 0. 4086 0.2102 18 12.0039 1. 5051 0.7785 01 0.6768 0. 0846 0.0421 03 1036.5000 203.9650 143.2050 06 39.7750 7. 9550 5.7700 2 8 3.9950 1.0200 0.7950 05 2.6954 0. 5430 0.3551 18 0.3607 0. 0508 0.0310 26 2.3150 0. 5050 0.3650 29 1.8300 0. 2352 0. 1520 12 1.5361 0. 1964 0.1146 14 14.0000 1.9850 1.3550 07 21. 7600 3.3450 2.3150 08 154.9600 15.2900 10.1 100 06 34.9750 6.3100 3.1750 19 1.2517 0. 1280 0.0739 29 3. 3673 0. 4505 0.2928 19 1.4 92 8 0. 1998 0.1199 08 18.5050 2. 1300 1 .9300 08 102.9800 11. 1450 7.1450 07 35.4350 4. 8450 3. 2800 12 1.1846 0. 1553 0.0865 19 3.7344 0.4829 0.2993 08 52 .7950 5.8350 3.8250 12 2.1478 0.2111 0.1122 03 12.6550 1.970 0 0. 6700 19 1.8483 0.2728 0.1511 08 177.6650 6.3100 3.1750 07 12.9500 1. 0600 0.5850 06 104.7050 20.5750 16.0650 03 66.6900 15.4900 9.4050 03 8607.0000 1708.7400 1261.3799 12 10.4262 1. 2318 0.6441 18 2.1294 0. 2808 0.1809 01 5.9601 0. 8388 0.4581 26 2.1747 0.5115 0.3 060 28 8.9331 2.0253 1.2237 03 78.3000 15.0300 1 1.2 250 12 7.2684 0. 7425 0.4899 19 2.3463 0. 4283 0.3075 08 0.4096 • 0.0615 0.0410 07 37.5000 5. 1500 3.3800 18 0.1538 0. 0265 0.0132 08 15.4350 1. 7 95 0 1.2950 19 2.9700 0.4900 0.2500 06 4.1600 0.8750 0.5600 19 13. 2450 1.8850 1.1050 08 119.1200 12.8800 8.4250 07 13.3850 1.5500 1.0450 03 0.2005 0.0993 0.0765 26 0.0 84 1 0. 0299 0.0157 18 1.2598 0. 1914 0.0651 12 1. 7927 0. 2972 0.1431 08 159.4600 16.6100 9.4 350 07 0.9458 0. 1760 0.1261 - 141 -1 16 03/08/76 1 65 90- 100 08 102.8700 10.6850 6. 6050 117 03/08/76 1 65 90- 100 05 0.0289 0. 0115 0.0072 118 27/07/76 1 95 20 -30 03 5.5710 1.1911 0.8128 1 19 27/07/76 1 95 20 -30 19 5. 1550 0.6734 0.4718 120 27/07/76 1 95 20 -30 12 2.8377 0. 3288 0.1991 121 27/07/76 1 95 20 -30 08 6.4184 0.6869 0.4476 122 27/07/76 1 95 20 -30 18 5. 1469 0.5896 0.3410 123 27/07/76 1 95 30 -40 03 101.5150 18.8700 15.2200 124 27/07/76 1 95 30 -40 06 7.7150 1. 3300 0.9150 125 27/07/76 1 95 30 -40 18 12.1050 1. 1250 0.7850 126 27/07/76 1 95 30 -40 08 25.7850 2. 1600 1.4800 127 27/07/76 1 95 30 -40 19 8.2500 1.3100 1.0850 128 27/07/76 1 95 30 -40 12 16.7500 1.9050 1.5950 129 27/07/76 1 95 40 -50 08 570.7849 47.2100 33.8850 130 27/07/76 1 95 40 -50 07 26.5200 2.0750 1. 5150 131 27/07/76 1 95 40 -50 19 25.8300 3. 1900 1.8350 132 27/07/76 1 95 40 -50 06 37.8100 5.3700 4.0650 133 27/07/76 1 95 40 -50 01 2.6062 0.3195 0. 1724 134 27/07/76 1 95 40 -50 18 1.5821 0. 1304 0.0890 135 27/07/76 1 95 40 -50 21 2. 1 989 0. 2472 0.1563 136 27/07/76 1 95 40 -50 12 11.4400 1.2000 1.0050 137 27/07/76 1 95 40 -50 03 7.6900 1. 2900 1.0150 138 27/07/76 1 95 50 -60 08 108.6100 13.7900 9.1550 139 27/07/76 1 95 50 -60 07 31.0800 5. 5550 3.6750 140 27/07/76 1 95 50 -60 19 52.5500 18. 7700 15.6050 141 27/07/76 1 95 50 -60 12 8.5800 1.5450 1.2200 142 27/07/76 1 95 50 -60 21 9.0000 1. 1600 0.9600 143 27/07/76 1 95 50 -60 16 0. 9015 0. 1039 0.0568 144 27/07/76 1 95 60 -70 08 2 05.9 95 0 21. 1850 13.9850 145 27/07/76 1 95 60 -70 07 71.4000 10.3550 7.2100 146 27/07/76 1 95 60 -70 12 23.3 565 0. 3875 0.2426 147 27/07/76 1 95 60 -70 19 9.5694 1. 0723 0. 5183 148 27/07/76 1 95 70 -80 08 8.4700 0.9550 0.7700 149 2 7/07/76 1 95 70 -80 07 34.9300 5.2000 3. 4250 150 27/07/76 1 95 70 -80 32 14.1 950 2. 1800 1.5650 151 27/0 7/76 1 95 80 -90 03 5.2741 1.0257 0.7345 152 27/07/76 1 95 80 -90 19 1.7023 0.2125 0.1396 153 27/07/76 1 95 80 -90 08 29.2250 3. 1600 2.2800 154 27/07/76 1 95 8 0 -90 07 108.9700 15.7850 9.9700 155 27/07/76 1 95 90-100 08 146.2150 20.2550 14.5650 156 27/07/76 1 95 90- 100 07 32.4750 2. 8300 2.1000 - 142 -ASH-FREE LOCATION SPECIES WEIGHT WEIGHT DRY WEIGHT DATE QUADRAT (G/io/r) (G/10M2) (G/10M2) 1 10/1 1/75 2 00 40- 50 15 So.3100 13.0600 2 10/1 1/75 2 00 40- 50 19 26.1500 3.5400 1.7950 3 io/r 1/75 2 00 40- 50 21 1.8549 0. 2727 0.1930 4 10/1 1/75 2 00 40- 50 27 0.6478 0. 1043 0.0568 5 10/1 1/75 2 00 40- 50 16 0.5302 0.C886 0.0579 6 10/1 1/75 2 00 40- 50 12 9.1300 1.5100 1.1500 7 10/1 1/75 2 00 50- 60 19 590.7000 8 1. 6 20 0 43.2650 8 10/1 1/75 2 00 50- 60 28 33.1100 6. 4 100 3. 2950 9 10/1 1/75 2 00 50- 60 15 90.9900 11 . 4200 6.9050 10 10/1 1/75 2 00 50- 60 1 1 166.4399 26.8500 14.2150 11 10/1 ' 1/75 2 00 50- 60 26 29.7000 4. 0100 2.7850 12 10/1 1/75 2 00 50- 60 23 11.0400 1.2938 0.8025 13 10/1 1/75 2 00 50- 60 03 34.9100 8.5600 6.5050 14 10/1 1/75 2 00 50- 60 13 9.0900 2. 0218 1.3464 15 10/1 1/75 2 00 50- 60 08 19.1500 2. 1899 1.3032 16 10/1 1/75 2 00 50- 60 12 9.8900 1. 1897 0.8325 17 io/r 1/75 2 00 50- 60 21 18. 0400 2.1508 1.5943 18 10/1' 1/75 2 00 50- 60 01 0.0839 0. 0108 0.0071 19 10/1' 1/75 2 00 50- 60 16 0.5793 0.0702 0.0450 20 10/1 1/75 2 00 50- 60 18 2.3471 0.2741 0.1905 21 10/1' 1/75 2 00 50- 60 05 0.3337 0. 0666 0.0414 22 10/1 1/75 2 00 50- 60 07 10.0801 1.9792 0.9223 23 10/1 1/75 2 00 50- 60 06 0.5 20 5 0.1276 0.0S18 24 io/r 1/75 2 00 60- 70 27 0.1726 0. 0654 0.0313 25 10/1 ' 1/75 2 00 60- 70 19 616.5100 92.7200 48.9200 26 10/1" 1/75 2 00 60- 70 15 302.3599 41.3500 24.7450 27 10/1' 1/75 2 00 60- 70 06 58.1200 12.8300 9.3200 28 10/1 1/75 2 00 60- 70 26 22.5000 4. 2200 2.7400 29 10/r 1/75 2 00 60- 70 12 5.9400 1.0600 0.7250 30 10/1 1/75 2 00 60- 70 11 60.5000 20.3900 1 1.9400 31 10/1 1/75 2 00 60- 70 08 31.9000 2. 7600 1.6600 32 10/1 1/75 2 00 60- 70 05 3.2400 0.8537 0.5842 33 10/T 1/75 2 00 60- 70 18 3.3416 0.4581 0.3434 34 10/1 1/75 2 00 60- 70 23 0.6200 0. 1018 0.0639 35 10/1 1/75 2 00 70- 80 19 15020.6500 2268.9805 1151.5C50 36 10/1 1/75 2 00 70- 80 15 781 .5999 87. 1000 49.5400 37 10/1 1/75 2 00 70- 80 12 111.5000 18.7000 12.5550 38 10/1 1/75 2 00 70- 80 13 10.8000 3.7795 2.6450 39 10/1 1/75 2 00 70- 80 26 37.7500 5. 2895 3.8552 40 10/1 1/75 2 00 70- 80 05 5.3965 1. 1315 0.8472 41 10/1 1/75 2 00 70- 80 01 1.6910 0.2190 0. 1560 42 10/1 1/75 2 00 70- 80 28 29.6500 8.1000 4.1550 43 10/1 1/75 2 00 70- 80 18 1.7778 0. 2200 0.1478 44 10/1 1/75 2 00 70- 80 27 8.1455 1.4740 0.8067 45 10/1 1/75 2 00 70- 80 17 57.3500 7.6000 4.3700 46 10/1 1/75 2 00 70- 80 29 21.3720 2.2630 1.5358 47 10/1 1/75 2 00 70- 80 30 9.1785 1.5230 0.9002 48 10/1 1/75 2 00 80- 90 15 46.8200 7.1250 4.1800 49 10/1 1/75 2 00 80- 90 19 1306.7649 200.6600 104.2400 50 10/1 1/75 2 00 80- 90 06 26.3450 9.6150 7.2650 51 10/1 1/75 2 00 80- 90 1 1 24.4850 5.0300 3.2100 52 10/1 1/75 2 00 80- 90 07 41.5800 2. 7650 1.6550 53 10/1 1/75 2 00 80- 90 . 08 16.2400 1.8950 1. 1250 54 10/1 1/75 2 00 80- 90 12 31.9700 5.8300 3.9100 55 10/1 1/75 2 00 80- 90 03 2.5750 0.6550 0.5300 - 143 -56 10/11/75 2 00 80 -90 26 15.1850 2. 9 150 1. 8100 57 10/11/75 2 00 80 -90 27 0.2763 0. 0824 0. 4553 58 10/11/75 2 00 80 -90 23 0.7448 0. 1144 0. 0730 59 10/11/75 2 00 80 -90 16 2.9408 0. 4470 0. 2853 60 10/11/75 2 00 80 -90 01 0.1499 0. 0324 0. 0215 61 10/11/75 2 00 90- 100 19 400.4800 64. 5100 35. 5350 62 10/11/75 2 00 90- 100 03 3.8900 1. 0567 0. 7789 63 10/11/75 2 00 90- 100 15 4.1800 0. 5678 0. 3295 64 10/11/75 2 00 90- 100 18 1.3172 0. 1758 0. 0668 65 10/11/75 2 00 90- 100 01 0.0603 0. 0095 0. 0069 66 10/11/75 2 00 90- 100 06 0.2573 0. 0366 0. 0263 67 10/11/75 2 00 90-100 16 0.4718 0. 0538 0. 0344 68 10/11/75 2 00 90- 100 17 7.3291 0. 6472 0.3303 69 10/11/75 2 00 90- 100 11 0.3866 0. 1014 0. 0587 70 10/11/75 2 00 90- 100 21 1.6267 0. 212 1 0.1379 71 10/11/75 2 00 90- 100 31 0.2648 0. 0762 0. 0388 72 10/11/75 2 00 90- 100 07 1.9939 0. 2807 0. 1322 73 10/11/75 2 00 90- 100 12 0.6059 0. 0777 0. 0549 - 144 -APPENDIX II A) Numerical species code for faunal assessment data in Appendix II (B). B) Faunal assessment data for seasonal collections at 95 m within Site 1. A. 01 Mytilus edulis 02 Amphithoe sp. 03 Notoacmea scutum 04 Margarites pupillus (parental) 05 Margarites pupillus (juvenile) 06 Strongylocentrotus droebachiensis 07 Lacuna marmorata 08 Mitrella gouldii 09 Tonicella liniata 10 Gnorimosphaeroma oregonense Dana 11 Idotea wosnesenskii Brandt 12 Unidentified polycheate 13 Pugettia richii 14 Amphilochus sp. 15 Metacaprella anomala 16 Alvinia spp. 17 Pandora sp. 18 Strongylocentrotus purpuratus Stimpson 19 Disporella sp. 20 Ocenebra sp. 21 Acmaea mitra 22 Cancer oregonensis 2 3 Odostomia spp. 24 Hiatella arctica 25 Granulina margaritula 26 Balcis mi cans 27 Bittium eschrichtii 28 Lirularia lirulata 29 Chi amys hastatus 30 Cancer branneri Rathbun 31 Nereis pelagica 32 Pagurus kennerlyi 33 Hemigrapsus nudus 34 Clinocardium sp. 35 Anatanias normani Richardson 36 Crepipatella lingulata Gould 37 Leptosynapta clarki Heding 38 Searlesia dira Reeve 39 Hyas lyratus Dana - 145 -B. WET DRY SPECIES WEIGHT WEIGHT DATE QUADRAT N/M (G/M2) (.G/M2) 1 25/05/76 30 04 1744 74. 7920 4 8. 539 2 2 25/05/76 30 06 32 50.3296 21.2704 3 25/0 5/76 30 01 48 0.3216 0. 2912 4 25/05/76 30 07 336 5.0624 4. 0352 5 25/05/76 30 08 128 8.7632 6. 7760 6 25/05/76 30 09 16 15.0528 9. 0064 7 2 5/0 5/76 30 02 384 11.3696 1. 9808 8 25/05/76 30 10 64 0.6432 0.244 8 9 25/05/76 30 1 1 16 0.5712 0. 2448 10 25/05/76 30 12 16 0.3072 0. 1424 11 25/05/76 30 13 32 6.0544 1. 6736 12 25/05/76 40 04 73 6 8.7520 4. 1920 13 25/05/76 40 09 144 9.8592 3.3936 14 25/05/76 40 02 16 0.9824 0. 1456 15 25/05/76 40 07 16 0.3008 0. 187 2 16 25/05/76 40 08 32 2.2448 1.3536 17 25/05/76 40 01 48 1.2480 0. 4576 18 25/0 5/76 40 14 48 0.2416 0.0400 19 25/05/76 40 27 16 0.7040 0. 5408 20 25/05/76 40 16 64 0.2592 0. 2384 21 25/05/76 40 17 32 1. 3008 0. 6880 22 25/05/76 50 04 640 23.0720 13.8928 23 25/05/76 50 06 16 51.5792 1 6. 3488 24 25/05/76 50 01 16 0.1760 0. 1744 25 25/05/76 50 09 96 3.8592 2. 4976 26 25/0 5/76 50 08 48 1.9360 1. 4672 27 25/05/76 50 07 16 0.2640 0. 1808 28 25/05/76 50 18 16 15.2992 8.0816 29 25/05/76 50 19 32 2.6224 0. 5712 30 25/05/76 50 20 48 1.0960 0. 6368 31 25/05/76 60 21 16 108.5984 81.137 6 32 25/05/76 60 17 16 1 .2720 0. 7936 33 25/05/76 60 03 16 0. 5952 0.3296 34 25/05/76 60 16 16 0.0208 0. 0064 35 25/05/76 70 01 608 4.7440 2. 3152 36 25/05/76 70 07 16 0.3136 0. 1376 37 25/05/76 70 09 64 10.0368 4.2112 38 25/0 5/76 70 04 64 0.4912 0. 3040 39 25/0 5/76 70 16 32 0.1104 0. 0768 40 25/05/76 70 23 32 0.2688 0. 0912 41 25/05/76 70 03 16 1. 0912 0. 4656 42 25/05/76 70 24 16 0.4080 0. 1408 43 25/05/76 80 13 16 0.8288 0. 3328 44 25/05/76 80 17 16 0.7472 0.5712 45 2 5/0 5/76 80 03 48 7.1456 3.4768 46 25/05/76 80 09 16 16.5360 8. 7424 47 25/0 5/76 80 04 48 2.1504 0. 7872 48 25/05/76 80 23 16 0.0752 0. 0592 49 25/05/76 80 16 32 0.0784 0. 0544 50 25/05/76 80 25 16 0.0656 0.046 4 51 25/05/76 80 26 16 0.0896 0. 056 0 52 25/05/76 90 09 192 95.3872 4 1. 3648 53 25/0 5/76 90 01 848 15.4192 4.2960 54 25/05/76 90 24 16 0.8448 0. 3232 55 25/05/76 100 09 64 34.8784 18. 5936 56 25/05/76 100 01 128 2.4640 1. 2656 57 25/05/76 100 28 16 0.4208 0. 2144 58 25/05/76 100 23 32 0.0960 0. 0560 59 25/05/76 100 26 16 0.1536 0. 1280 60 14/06/76 30 13 96 46.2352 16. 4960 61 14/0 6/76 30 08 400 20.4720 1 4. 2064 62 14/06/76 30 27 192 19.4672 16.2656 63 14/06/76 30 19 64 20.3312 8. 0240 6a 14/06/76 30 01 64 0.4560 0.4400 65 14/06/76 30 20 80 13.5344 10. 6256 66 14/06/76 30 17 16 0.4640 0.4592 67 14/0 6/76 30 09 32 4.9920 3.6784 68 14/06/76 30 07 96 1.7424 1.3664 69 14/06/76 30 25 192 0. 6784 0. 5392 70 14/06/76 30 29 32 0.6304 0. 2768 71 14/0 6/76 30 04 2112 49.9344 26. 0752 72 14/06/76 30 16 112 0.0496 0.0464 73 14/06/76 30 31 16 1.0048 0. 0608 74 14/06/76 30 02 256 0.6064 0. 1456 75 14/06/76 30 14 48 0.4960 0. 1024 76 14/06/76 30 32 48 1.8624 0.3712 77 14/06/76 30 23 80 0.2304 0.2016 78 14/06/76 40 04 1264 53.1856 34.3632 79 14/06/76 40 06 64 37.1632 52. 2160 80 14/06/76 40 13 96 58.1264 1 8. 9584 81 14/06/76 40 27 64 5.2336 4.4192 82 14/06/76 40 02 240 11.2000 2. 8640 83 14/06/76 40 31 32 8.3056 2.1440 84 14/06/76 40 01 96 3.0976 1. 9424 85 14/06/76 40 09 48 8.9104 6. 0048 86 14/06/76 40 07 144 3.1280 2. 1504 87 14/0 6/76 40 08 96 6.4160 4. 6064 88 14/06/76 40 29 32 0.5360 0. 4784 89 14/06/76 40 33 16 2. 7008 0. 1952 90 14/06/76 4 0 34 48 8.0576 4. 5248 91 14/06/76 50 01 1728 45.4352 1 5. 4976 92 14/06/76 50 04 432 16.1040 27.7840 93 14/06/76 50 02 48 1.7104 0. 403 2 94 14/06/76 50 14 32 0.2496 0. 0400 95 14/06/76 50 09 80 1 1. 8240 6.1936 96 14/06/76 50 27 48 1 .5856 1. 3024 97 14/06/76 50 24 96 2.2208 0. 9856 98 14/06/76 50 20 16 0.9648 0. 7664 99 14/06/76 50 03 16 2.8400 1.7952 100 14/06/76 50 25 16 0.0784 0. 0560 101 14/06/76 50 31 48 2.2560 0. 2832 102 14/06/76 60 09 160 20.7024 1 0. 2160 103 14/06/76 60 17 16 8.5616 5. 0768 104 14/06/76 60 04 48 1.1328 0. 7376 105 14/06/76 60 24 32 0.8640 0.4016 106 14/06/76 60 32 16 1.0064 0. 1360 107 14/06/76 60 16 16 0.0352 0. 022 4 108 14/06/76 60 07 16 0.9856 0. 6656 109 14/06/76 60 25 32 0.1568 0. 0832 1 10 14/06/76 60 01 15968 1047.8113 313.5952 111 14/06/76 60 26 32 0.1920 0. 0528 112 14/06/76 60 03 16 0.6736 0. 3280 1 13 14/06/76 70 01 5648 293.9121 129.1760 114 14/06/76 70 21 16 18.5600 1 2. 6832 1 15 14/06/76 70 09 64 71. 2096 39.2668 116 14/06/76 70 25 1 17 14/06/76 70 03 118 14/06/76 80 01 1 19 14/06/76 80 09 120 14/0 6/76 80 08 121 14/06/76 80 16 122 14/06/76 80 28 123 14/06/76 90 09 124 14/06/76 90 01 125 14/06/76 90 20 126 14/0 6/76 90 03 127 14/06/76 90 25 128 14/06/76 90 28 129 14/06/76 90 35 130 14/06/76 100 31 131 14/06/76 100 14 132 14/06/76 100 02 133 14/06/76 100 01 134 14/06/76 1 00 18 135 14/06/76 100 20 136 14/06/76 100 09 137 14/06/76 100 08 138 14/06/76 1 00 36 139 08/07/76 30 09 140 08/07/76 30 08 141 08/07/76 30 04 142 08/07/76 30 21 143 08/07/76 30 25 144 08/07/76 30 14 145 08/07/76 30 23 146 08/07/76 30 26 147 08/07/76 30 13 148 08/07/76 30 02 149 08/07/76 30 27 150 08/07/76 30 31 151 08/07/76 30 07 152 08/07/76 30 01 153 08/07/76 30 34 154 08/07/76 40 37 155 08/07/76 40 13 156 08/07/76 40 01 157 08/07/76 40 24 158 08/07/76 40 34 159 08/07/76 40 07 160 08/07/76 40 20 161 08/07/76 40 25 162 08/07/76 40 04 163 08/07/76 40 27 164 08/07/76 40 08 165 08/07/76 40 32 166 08/07/76 40 29 167 08/07/76 40 02 168 08/07/76 50 25 169 08/07/76 50 23 170 08/07/76 50 02 171 08/07/76 50 14 172 08/07/76 50 09 173 08/07/76 50 07 174 08/07/76 50 04 175 08/07/76 50 28 - 147 -16 0.0608 0. 0240 16 34. 81 12 23.1152 112 1.7824 0. 9040 32 3.4384 2. 2128 16 0.8400 0. 6336 32 0.0752 0. 0528 80 0.6256 0. 3872 128 49.4336 26.8400 2832 84.0800 33.0752 32 35.3648 25. 7392 48 16.2544 9. 6464 256 0.9808 0. 6704 16 0.0656 0. 0400 48 0.1952 0. 0608 16 1.3696 0. 0688 32 0.0512 0.0368 48 0.1072 0.0336 16 0.0800 0. 0352 16 3.0864 2. 1232 32 8.0512 5. 9120 48 6.8960 3.0128 64 2.1040 1.4880 16 1.4592 0. 9072 112 65.0576 36.7824 96 6.9744 5. 139 2 2048 86.1360 53. 0400 16 0.3680 0. 3232 80 0.36 16 0. 2496 32 0.1792 0. 0432 16 0.0736 0. 0560 16 0.0208 0.0112 16 1.8592 0. 3696 32 0.4896 0. 107 2 16 1. 0416 0. 8528 16 0.2512 0. 1296 54 4 3.8736 2. 5424 32 8.4880 4. 7728 16 0.5328 0. 3984 16 126.2608 7. 9184 48 85. 2464 1 8. 0976 16 18.9696 9. 0544 32 1. 2384 0. 7296 48 1.2448 0. 931 2 288 3.1168 1. 8688 32 4.8912 3.7920 32 0.1376 0.0784 768 26.5008 14.5776 304 39.3072 31. 0624 416 25.4128 17.1744 64 3.5104 1. 1184 32 0.4688 0. 3024 16 0.5136 0. 1440 16 0.1056 0. 0640 32 0.1072 0. 0720 16 0.1184 0. 0288 32 0.2224 0. 0544 32 3.2336 1. 6320 176 1.8496 0. 9504 496 23.6528 12.6944 112 5.2784 2. 8544 - -176 08/07/76 50 13 32 50.4272 1 1. 5120 177 08/07/76 50 33 16 5.6192 1.2320 178 08/07/76 50 01 48 0.7936 0.3664 179 08/07/76 50 19 48 11.9664 3. 1264 180 08/07/76 50 08 16 0.8400 0. 5888 181 08/07/76 60 09 96 67.9712 35. 321 6 182 08/07/76 60 07 80 1.5200 0. 8352 183 08/07/76 60 21 32 20.5600 15.0272 184 08/07/76 60 25 16 0.0976 0. 0608 185 08/07/76 60 08 16 0.7200 0. 5632 186 08/07/76 60 01 32 0.8272 0. 596 8 187 08/07/76 60 28 16 0.7776 0. 5024 188 08/07/76 60 20 16 0.4112 0. 3360 189 08/07/76 70 09 32 289.1968 133.9840 190 08/07/76 70 01 1296 93.9984 41.6960 191 08/07/76 70 25 48 0.2608 0. 1760 192 08/07/76 70 08 64 2.7376 1.8864 193 08/07/76 70 28 32 0.3536 0. 2176 194 08/07/76 80 09 32 62.1408 32. 3648 195 08/07/76 80 12 16 33.2800 0. 2720 196 08/07/76 80 21 16 112.5360 79.9024 197 08/07/76 80 25 16 0.0912 0.0736 198 08/07/76 80 01 112 2.5792 1.5712 199 08/07/76 80 07 16 1.5536 0. 4720 200 08/07/76 90 01 512 25.7472 12.8496 201 08/07/76 90 09 96 39.6304 21. 7296 202 08/07/76 90 21 16 16.6560 11.5264 203 08/07/76 90 25 32 0.2352 0. 1760 204 08/07/76 90 20 48 2.3056 1. 8544 205 08/07/76 90 33 16 0.2592 0. 1200 206 08/07/76 100 28 32 0.2032 0. 1504 207 08/07/76 100 02 48 0.1792 0. 0560 208 08/07/76 1 00 09 80 19.3760 1 0. 0704 209 08/07/76 100 25 272 1.1824 0. 7488 210 28/07/76 30 07 57088 175.6880 86.3280 211 28/07/76 30 04 640 32. 8736 18. 2752 212 28/07/76 30 08 176 12.9984 8. 7072 213 28/07/76 30 13 16 29.8256 6.9984 214 28/07/76 30 09 48 13.5872 6. 854 4 215 28/07/76 30 01 32 3.3152 1. 6688 216 28/07/76 30 29 16 0.6176 0. 3008 217 2 8/07/7 6 30 34 16 2.1728 1. 3952 218 28/07/76 30 27 32 4.3168 3.3664 219 28/07/76 40 25 32 0.2032 0. 1472 220 28/07/76 40 27 176 22.4224 17.3808 221 28/07/76 40 01 16 2.4400 0. 9456 222 28/07/76 40 09 80 13.4672 6. 6160 223 28/07/76 40 08 1 12 7.4816 4. 9728 224 28/07/76 40 02 16 0.1776 0. 0432 225 28/07/76 40 14 16 0.0896 0. 0576 226 28/07/76 40 29 32 0.7056 0. 4352 227 28/07/76 40 07 256 2.5216 1. 5376 228 28/07/76 40 13 32 41.3696 13. 1232 229 28/07/76 40 39 16 9.0400 1.9120 230 28/07/76 40 04 2064 89.1056 48. 844 8 231 28/07/76 50 01 128 3. 8032 . 0. 9872 232 28/07/76 50 24 48 1 .3872 0. 5600 233 28/07/76 50 13 32 8.3280 2. 4432 234 23/07/76 50 27 16 0.7456 0. 5536 235 28/07/76 50 14 16 0.0384 0. 017 6 236 28/07/76 50 08 16 0.9472 0.6272 237 28/07/76 50 28 96 3.4016 1. 8768 238 28/07/76 50 05 80 0.4448 0. 2256 239 28/07/76 50 09 32 1.3008 0. 673 6 240 28/07/76 50 16 32 0.0800 0. 0368 241 28/07/76 50 23 16 0.0368 0. 0192 242 28/07/76 50 07 48 0.6432 0.3936 243 28/07/76 50 26 16 0.0912 0. 0496 244 28/07/76 60 01 624 60.5008 19. 4144 245 28/07/76 60 20 48 2.4768 1.2976 246 28/07/76 60 09 32 16.1616 5.5120 247 28/07/76 60 03 32 1 1.8912 5. 2208 248 28/07/76 60 08 80 5.3120 3. 1232 249 28/07/76 70 01 3984 680.6001 312.9919 250 28/07/76 70 09 32 8.0448 3.2096 251 28/07/76 70 25 64 0.3760 0. 2128 252 28/07/76 70 07 16 0.7680 0. 5248 2 53 28/07/76 70 08 32 2.2256 1. 435 2 254 28/07/76 70 26 16 0.0912 0. 0272 255 28/07/76 80 21 16 55. 1056 34. 8400 256 28/07/76 80 09 16 24.7504 10.0736 257 28/07/76 80 25 16 0. 1024 0. 0640 258 28/07/76 80 16 16 0.0336 0. 0272 259 28/07/76 90 01 5888 951.0400 433. 9199 260 28/07/76 90 09 48 6.6208 2. 5856 261 28/07/76 90 20 48 4.8304 3. 2272 262 28/07/76 90 24 16 0.6192 0.2736 263 28/07/76 90 25 32 0.2144 0. 1184 264 28/07/76 1 00 09 48 9.4768 3.96 8 0 265 28/07/76 100 33 16 0.2240 0. 0480 266 28/07/76 1 00 22 16 36.9072 12.7424 267 28/07/76 100 16 16 0.0256 0. 0080 268 28/07/76 100 25 16 0. 1040 0. 0656 269 28/07/76 100 08 16 0.9200 0. 5888 270 28/07/76 1 00 20 32 6.3760 4. 0384 271 18/08/76 30 23 16 0.0848 0. 0656 272 18/08/76 30 07 11680 65.5968 24.6576 273 18/08/76 30 05 2048 4. 5472 2. 6832 274 18/08/76 30 04 1168 62.0896 35.0544 275 18/08/76 30 34 112 5.2224 3. 3408 276 18/08/76 30 08 64 4.2688 2. 9536 277 18/0 8/76 30 09 16 3.2416 1.8192 278 18/08/76 30 13 16 4.8176 1. 4304 279 18/0 8/7 6 30 24 16 3.2352 1. 4384 280 18/0 8/76 30 27 32 4.1568 3.2576 281 18/08/76 30 25 32 0.1776 0.1104 282 18/08/76 40 25 320 1.6000 1.0112 283 18/08/76 40 04 896 28.0512 15. 5920 284 18/08/76 40 05 2208 9.2720 4. 3888 285 18/08/76 40 15 32 0.1008 0. 0423 286 18/08/76 40 07 3008 1 5.7056 7. 5888 287 18/08/76 40 16 80 0. 1616 0. 0816 288 18/08/76 40 23 160 0.4032 0. 1872 289 18/0 8/76 40 08 16 1.2432 C. 9200 290 18/08/76 40 13 16 0.3120 0. 2416 291 18/03/76 40 22 32 0.4208 0. 1536 292 18/08/76 40 09 32 3.3232 1.4880 293 18/08/76 40 27 16 0.2608 0. 203 2 294 18/08/76 50 05 5920 37. 5568 1 9. 7600 295 18/08/76 50 07 368 1. 2928 0. 8320 - IbU -296 18/08/76 50 23 928 3.7280 2. 6880 297 18/08/76 50 16 688 1.8960 1.2128 298 18/08/76 50 15 32 0.1216 0. 0304 299 18/08/76 50 25 288 1.0528 0. 6560 300 18/08/76 50 06 32 0.6912 0. 3872 301 18/08/76 50 04 176 6. 9392 4.0704 302 18/0 8/76 50 28 528 7. 1216 4. 0272 303 18/08/76 50 27 448 17.2064 13. 7136 304 18/08/76 50 34 32 0.4432 0. 3424 305 18/08/76 50 20 80 2.1872 1. 6208 306 18/08/76 50 09 16 0.9040 0. 5648 307 18/08/76 50 08 496 10.6928 5. 8048 308 18/08/76 60 01 560 57. 0384 39. 0960 309 18/08/76 60 09 48 30.5504 1 5. 1232 310 18/08/76 60 03 16 0.3088 0. 3008 311 18/08/76 60 25 16 0.0640 0. 0384 312 18/08/76 60 23 32 0.1552 0. 1056 313 18/08/76 60 16 16 0.0304 0. 0144 314 18/08/76 70 01 912 193.1456 103.9456 315 18/08/76 70 09 32 7.2976 3.7664 316 1 8/08/76 70 33 16 8.9952 4. 3280 317 18/08/76 70 15 48 0.1712 0.0720 318 18/08/76 70 25 64 0.3312 0. 2160 319 18/03/76 70 23 16 0.0656 0. 0288 320 18/0 8/76 70 08 112 6.7600 4. 8432 321 18/0 8/76 70 16 16 0.0272 0. 0176 322 18/08/76 80 09 48 46.9456 22.8896 323 18/08/76 80 03 16 0.6192 0. 3872 324 18/08/76 80 08 48 1 .8448 1. 3008 325 18/0 8/76 80 20 32 1.0048 0. 784 0 326 18/08/76 80 25 48 0.2784 0. 1936 327 18/08/76 80 28 32 0.4112 0. 2464 328 18/08/76 90 0 1 256 10.7168 6.516 8 329 18/08/76 90 03 32 18.4032 1 1. 7728 330 18/08/76 90 09 32 8.3744 5. 0496 331. 18/08/76 90 20 48 0.9872 0. 7824 332 18/0 8/76 90 25 64 0.2912 0. 2192 333 18/08/76 90 20 16 1.8368 1. 3920 334 18/08/76 90 28 16 0.2768 0. 2032 335 18/08/76 100 28 128 2.0704 1. 3728 336 18/08/76 100 15 32 0.0816 0. 0272 337 18/08/76 100 08 144 7.1424 4. 9152 338 18/0 8/76 1 00 22 160 2.3824 0. 7984 339 18/08/76 1 00 25 272 1. 3600 0..9056 340 18/08/76 1 00 09 112 11.6592 6.435 2 341 18/08/76 100 20 48 1.9680 1.4816 342 18/0 8/76 1 00 13 16 2.5883 0. 8160 343 18/08/76 100 14 64 0.0976 0. 0352 344 12/09/76 30 05 1648 5.9104 3.7936 345 12/09/76 30 07 1 1760 62.9824 33.3792 346 12/09/76 30 23 96 0.5104 0. 3200 347 12/09/76 30 16 176 0.5376 0. 2832 348 12/09/76 30 13 16 14.5312 3. 6832 349 12/09/76 30 06 16 14. 4016 6.6304 350 12/09/76 30 34 16 0.8496 0. 5552 351 12/09/76 30 24 16 1.0144 0. 395 2 352 12/09/76 30 04 672 37.5696 22.0496 353 12/09/76 30 08 160 9.8400 7. 064 0 354 12/09/76 40 05 54"0 22.1040 16. 0640 355 12/09/76 40 07 9840 50.6720 33.7200 356 12/09/76 40 25 357 12/09/76 40 23 358 12/09/76 40 16 359 12/09/76 40 04 360 12/09/76 40 12 361 12/09/76 40 27 362 12/09/76 40 08 363 12/09/76 40 32 364 12/09/76 40 13 365 12/09/76 40 20 366 12/09/76 40 28 367 12/09/76 50 27 368 12/09/76 50 08 369 12/09/76 50 33 370 12/09/76 50 05 371 12/09/76 50 07 372 12/09/76 50 23 373 12/09/76 50 16 374 12/09/76 50 28 375 12/09/76 60 09 376 12/09/76 60 01 377 12/09/76 60 22 378 12/09/76 60 07 379 12/09/76 60 05 380 12/09/76 60 06 381 12/09/76 60 28 382 12/09/76 60 08 383 12/09/76 60 26 384 12/09/76 60 16 385 12/09/76 60 23 386 12/09/76 70 01 387 12/09/76 70 09 388 12/09/76 70 05 389 12/09/76 70 22 390 12/09/76 70 34 391 12/09/76 70 20 392 12/09/76 70 28 393 12/09/76 70 25 394 12/09/76 70 26 395 12/09/76 70 31 396 12/09/76 80 09 397 12/09/76 90 09 398 12/09/76 90 01 399 12/09/76 90 03 400 12/09/76 90 25 401 12/09/76 1 00 22 402 12/09/76 1 00 09 403 12/09/76 100 25 404 12/09/76 100 13 405 12/09/76 1 00 28 406 12/09/76 1 00 14 407 12/09/76 100 20 408 07/10/76 30 05 409 07/10/76 30 07 4 10 07/10/76 30 30 411 07/10/76 30 34 412 07/10/76 30 20 413 07/10/76 30 08 4 14 07/10/76 30 25 4 15 07/10/76 40 13 - 151 -800 3.0080 2. 3120 480 1.8560 1.6160 560 1.0240 0. 8880 1568 70.8944 39. 9840 16 2.4160 0.4336 128 18.5264 14. 9632 240 15.6912 10.5376 16 4.9376 2. 3600 16 9. 8672 2. 7264 16 0.5728 0. 4976 112 1.4336 0. 9824 48 2.5504 2.1616 16 1 .1360 0.9072 16 0.3520 0. 2656 5472 29.5776 20. 1536 2192 8.2304 6. 2720 64 0.4272 0. 2816 27 2 0.5856 0. 5040 96 1.1488 0. 836 8 16 1.6224 1. 3680 352 50.8896 29. 939 2 32 0.5088 0. 3872 32 0.7232 0. 4128 320 0.9552 0.7312 16 0.1984 0. 1824 16 0.4144 0. 3008 16 0.9088 0. 684 8 16 0.0848 0. 0576 32 0.0896 0. 0672 48 0.1088 0. 0848 608 85.0928 34.4528 16 3. 1792 2. 0640 80 0.2768 0.219 2 32 0.2736 0. 2080 16 0.4096 0.3632 16 0. 1776 0. 1264 16 0.4832 0. 3248 160 0.7664 0. 6096 96 0.6128 0. 4704 16 0.2048 0. 1264 16 19.5952 12. 9728 80 14.5488 10.4736 368 32.1248 23. 2128 32 25.2224 16. 9328 80 0.3408 0. 2624 32 0.2784 0. 2320 48 7.8256 5.5136 . 176 0.7904 0. 6384 16 0.5328 0. 4336 16 0.3552 0.2592 80 0.0800 0. 0496 16 2.0464 1. 72C0 1 488 11.2128 7.4160 3072 35. 5760 2 1. 0624 16 2.2464 0. 8032 32 0.7920 0. 6544 16 11.1072 8.9712 112 7.9808 6. 004 8 144 0.6432 0. 4544 16 38.8848 1 2. 0320 416 07/10/76 40 30 48 18.9936 7. 5088 417 07/10/76 40 06 16 34.2896 17. 3856 418 07/10/76 40 09 16 0.9008 0. 6960 419 07/10/76 40 34 160 7.1328 5. 5264 420 07/10/76 40 27 128 17.2544 1 4. 6784 421 07/10/76 40 08 320 21.2240 16.2288 422 07/10/76 40 01 32 0.7584 0. 4400 423 07/10/76 40 25 144 0.5744 0. 4256 424 07/10/76 40 07 1648 20.5664 13.3760 425 07/10/76 40 05 2480 25. 8608 16.1504 426 07/10/76 40 04 176 8.2272 5.7280 427 07/10/76 40 23 32 0. 1344 0. 0928 428 07/10/76 40 16 16 0. 0624 0. 0224 429 07/10/76 50 25 464 2.3104 1.2464 430 07/10/76 50 28 304 5.0768 3. 2384 431 07/10/76 50 05 20064 174.7696 100.8064 432 07/10/76 50 07 5776 22.9680 15.2000 433 07/10/76 50 23 464 2.8576 2. 1888 434 07/10/76 50 16 8976 18.6352 13.0112 435 07/10/76 50 04 80 5.6576 3. 4848 436 07/10/76 50 27 160 9.1904 7. 4688 437 07/10/76 50 20 32 2. 1 152 1. 6080 438 07/10/76 50 34 16 1.2976 0. 9056 439 07/10/76 50 30 16 1.0704 0. 3648 440 07/10/76 50 09 16 5.5120 2. 9552 441 07/10/76 50 32 16 0.2448 0.108 8 442 07/10/76 60 32 16 0. 1248 0.0912 443 07/10/76 60 05 64 0.3120 0. 2256 444 07/10/76 60 0 1 112 0.5376 0. 3584 445 07/10/76 60 07 48 0.4640 0. 3088 446 07/10/76 70 09 16 13.9712 8. 7120 447 0 7/10/76 70 08 80 4.5664 3. 4224 448 07/10/76 70 28 48 0.6208 0.5088 449 07/10/76 70 01 704 53. 5872 36. 8704 450 07/10/76 70 20 16 0.7312 0. 5312 45 1 07/10/76 70 30 32 0.8192 0. 5712 452 07/10/76 80 01 16 0.2208 0. 0992 453 07/10/76 80 05 64 1.4656 0. 9264 454 07/10/76 80 09 16 2.4128 1. 1328 455 07/10/76 80 20 32 0.6960 0.4992 456 07/10/76 80 32 16 0.1600 0. 0944 4 57 07/10/76 80 25 32 0.1712 0. 0992 458 07/10/76 80 08 16 0.7424 0. 5088 459 07/10/76 80 28 112 1. 9696 1. 2480 460 07/10/76 80 16 128 0.2352 0. 1664 461 07/10/76 80 23 64 0.3856 0. 2960 462 07/10/76 90 09 64 16.5152 10. 0528 463 07/10/76 90 0 1 112 0.7088 0. 6144 464 07/10/76 90 12 16 0.5856 0.5040 465 07/10/76 90 03 16 11.5568 7.3088 466 07/10/76 90 02 32 0.0352 0. 0160 467 07/10/76 1 00 09 32 7.1264 3. 0128 468 07/10/76 1 00 38 64 49.2656 34. 3504 469 07/10/76 1 00 20 32 1.1936 0. 9504 470 07/10/76 1 00 08 80 2.7808 2. 0432 471 07/10/76 100 24 16 0.4592 0.4192 472 07/10/76 100 25 384 2.0944 1. 3008 473 07/10/76 100 07 64 0.2288 0. 1920 474 07/10/76 1 00 26 16 0.2480 0. 1952 475 07/10/76 100 28 16 0.3200 0. 2368 - 153 -APPENDIX III Detritus assessment data for seasonal collections at 95 m within Site 1. - 154 -DRY ASH-FREE WEIGHT DRY WEIGHT DATE QUADRAT ( G/ti2 ) ( G/M2 ) 1 28/05/76 20 0. 13 0. 07 2 28/05/76 30 0. 62 0. 12 3 28/05/76 40 1. 15 0. 15 4 28/05/76 50 1. 01 0. 19 5 28/05/76 60 0. 60 0. 15 6 28/05/76 70 1. 39 0. 28 7 28/05/76 80 1. 61 0. 33 8 28/05/76 90 1. 75 0. 27 9 28/05/76 100 1. 54 0. 27 10 17/06/76 20 0. 25 0. 11 11 17/06/76 30 1. 19 0. 32 12 17/06/76 40 0. 87 0. 30 13 17/06/76 50 0. 86 0. 27 14 17/06/76 60 1. 01 0. 32 15 17/06/76 70 1. 17 0. 43 16 17/06/76 80 0. 76 0. 15 17 17/06/76 90 1. 23 0. 24 18 17/06/76 100 1. 48 0. 37 19 08/07/76 20 0. 32 0. 14 20 08/07/76 30 2. 20 0. 55 21 08/07/76 40 4. 46 0. 43 22 08/07/76 50 4. 13 0. 76 23 08/07/76 60 2. 11 0. 61 24 08/07/76 70 3. 97 0. 74 25 08/07/76 80 2. 52 0. 60 26 08/07/76 90 3. 63 0. 58 27 08/07/76 100 2. 98 0. 54 28 29/07/76 20 0. 28 0. 12 29 29/07/76 30 2. 99 0.72 30 29/07/76 40 2. 87 0. 48 31 29/07/76 50 1. 98 0. 50 32 29/07/76 60 2. 06 0. 57 33 29/07/76 70 1. 66 0. 46 34 29/07/76 80 2. 19 0. 54 35 29/07/76 90 2. 17 0. 50 36 29/07/76 100 2. 24 0. 54 37 20/08/76 20 0. 31 0.16 38 20/08/76 30 5. 38 1. 11 39 20/08/76 40 6. 60 1.39 40 20/08/76 50 6. 00 1. 20 41 20/08/76 60 1. 57 0.42 42 20/08/76 70 3. 35 0. 69 43 20/08/76 80 0. 58 0. 13 44 20/08/76 90 1. 17 0. 26 45 20/08/76 100 1. 05 0. 23 46 12/09/76 20 0. 45 0. 16 47 12/09/76 30 1. 48 0. 32 48 12/09/76 40 0. 61 0. 13 49 12/09/76 50 1. 02 0. 19 50 12/09/76 60 0. 61 0. 10 51 12/09/76 70 0. 78 0. 21 52 12/09/76 80 0. 49 0. 09 53 12/09/76 90 0. 99 0.20 54 12/09/76 100 0. 82 0. 20 55 07/10/76 20 0. 29 0. 10 - lbb -56 07/10/76 30 0. 52 0. 12 57 07/10/76 40 0. 97 0. 28 58 07/10/76 50 1. 91 0. 55 59 07/10/76 60 0. 93 0. 30 60 07/10/76 70 0. 72 0. 23 61 07/10/76 80 0. 78 0. 34 62 07/10/76 90 0. 61 0. 22 63 07/10/76 100 1 . 75 0. 45 - 156 -APPENDIX IV Depth data (m below mean sea level) for the transects at 5, 35, 65 and 95 m within Site 1. Transect location Distance along transect (m) 00 05 10 15 20 25 30 35 40 45 50 55 60 65 70 75 80 85 90 95 100 05 m -1.4 -0.6 0.5 0.8 2.3 2.9 2.3 2.6 2.9 3.4 3.8 4.1 4.4 4.9 5.0 5.5 6.3 6.7 7.2 7.6 8.1 35 m -1.2 -0.6 0.3 1.2 2.1 2.3 2.4 2.4 2.6 2.7 3.7 4.3 4.9 5.5 6.1 6.4 7.0 7.9 8.5 9.1 9.8 65 m -1.1 -0.8 -0.2 0.8 2.0 1.8 1.3 1.5 1.8 2 . 3 2.9 3.7 4.4 4.7 5.3 6.1 6.9 7.5 8.1 9.0 9.6 95 m -0.9 -0.3 0.6 1.2 1.5 2.1 2.7 3.7 4.1 4.9 5.5 5.5 6.1 6.4 6.4 6.7 6.7 7.0 7.3 7.6 7.9 - 157 -APPENDIX V Litter decomposition experimental data. Length of Percentage of Species incubation period (days) original dry weight Plocamium coccineum var. pacificum 0 100.00 10 65.26 16 42.50 24 28.22 Rhodomela larix 0 100.00 6 86.213 48.73 25 6.48 Odonthalia floccosa 0 100.00 8 55.35 19 34.51 33 9.23 Iridaea cordata 0 100.00 2 97.39 8 55.66 13 0.13 Gigartina papillata 0 100.00 10 38.516 16.79 24 2.72 Constantinea subulifera 0 100.00 6 62.314 45.20 29 11.57 Fucus distichus 0 100.00 6 61.08 13 39.99 19 44.38 44 8.67 Nereocystis luetkeana (stipe) 0 100.00 6 29.713 5.38 19 0.01 Nereocystis luetkeana (lamina) 0 100.00 2 52.84 5.49 6 0.08 - 158 -Appendix V (continued) Laminaria saccharina 0 100.00 8 13.79 11.30 Laminaria groenlandica 0 100.00 3 30.14 8 11.16 9 10.13 - 159 -APPENDIX VI Oxygen consumed (mg) by microbes decomposing three particle sizes of the 10 detrital species in Experiment 1 following three periods of incubation. Species 5 days 10 days 20 days Plocamium coccineum var. pacificum 0.17 0.32 0.49 44-0 0.20 0.36 0.47 250-149 0.17 0. 33 0.47 1000-420 Rhodomela larix 0.22 0.31 0.42 0.16 0.29 0.45 0.25 0. 31 0.42 Odonthalia floccosa 0.19 0.37 0.46 0.15 0.28 0.49 0.19 0. 34 0.49 Iridaea cordata 0.50 0.57 0.65 0.42 0.45 0.64 0. 33 0.55 0.71 Gigartina papillata 0.26 0.28 0.38 0.20 0.33 0.42 0.22 0.30 0.36 Constantinea subulifera 0. 35 0.60 0.62 0. 37 0.54 0.67 0.35 0.52 0.64 Fucus distichus 0. 38 0.57 0.84 0.46 0.55 0.71 0. 39 0.63 0.85 Nereocystis luetkeana (stipe) 0.28 0. 36 0.52 0.29 0.35 0.50 0. 34 0.40 0.50 Nereocystis luetkeana (lamina) 0.29 0.36 0.57 0. 36 0.47 0.47 0.27 0.41 0.49 Laminaria saccharina 0.25 0.39 0.54 0.29 0. 37 0.46 0.29 0.32 0.47 Laminaria groenlandica 0. 31 0.35 0.45 0.25 0. 38 0.50 0.29 0.45 0.48 particle size -160 -APPENDIX VII Percentage of particulate material remaining following three periods of incubation for three particle sizes of the 10 detrital species decomposed in Experiment 2. Species 10 days 20 days 30 days Plocamium coccineum var. pacificum Rhodomela larix Odonthalia floccosa 100.0 114.8 94.9 98.1 102.6 107.7 111. 3 123.3 95.0 101.4 86.7 109.5 102.0 110.0 106.6 101.3 110.7 92.4 95.6 77.3 94.5 95.0 96.7 103.5 100.0 104.6 97.7 44-0 urn"1 250-149 urn| 1000-420 um; particle size Iridaea cordata Gigartina papillata 22.4 45.4 62.8 74.1 101.1 89.1 20.7 22.5 24.0 70.0 99.6 89.4 24.2 29.5 25.8 77.3 97.2 63.1 Constantinea subulifera 100.7 94.9 93.3 97.1 109.5 89.3 88.1 94.5 75.5 Fucus distichus 123.7 101.1 104.8 100.9 99.6 106.4 96.0 97.2 103.5 Nereocystis luetkeana (stipe) Nereocystis luetkeana (lamina) 54.2 70.9 81.2 63.9 65.0 66.7 44.4 52.2 60.6 51.2 47.4 59.6 48.2 35.0 62.2 49.2 49.1 37.1 Laminaria saccharina 73.6 77.2 81.4 71.1 70.1 67.2 72. 3 45.1 70.6 Laminaria groenlandica 60.4 62.8 68.6 67.1 59.6 63.6 72.5 55.1 61.8 - 161 -APPENDIX VIII FORTRAN G computer program for the simulation model of litter and detritus processing within Site 1. Main program: Accepts parameters determining the data to be processed, i.e. wet, dry or ash-free dry weight; sets the significance level of the chi-square test for patchiness in litter distribution; calls subroutines Ml, M2, M3 and M4. Ml: Creates a three dimensional matrix (species, quadrat, tran sect) of litter biomass data defining the areal distribution of litter within Site 1. The matrix is based on data from the transect collections at 5, 35, 65 m within Site 1 on 3 August and at 95 m on 27 July 1976. M2: Tests (chi-square) for patchiness in the distribution of specific litter within equivalent quadrats of the four tran sects defining the areal distribution of litter within Site 1. If the result is non-significant, the data are averaged to reduce the influence of sampling variability. M3: Calculates the equation (Figure 17) for the seasonal distri bution of total litter biomass within Site 1. M4: Performs the operations outlined in the flow chart in Figure 18. - 162 -1 INTEGER WTPAR 2 COMMON WTPAR /AREA1/ WTDAS(5,4,10) /AREA2/ DAY1(17), P(ll), REGWT( +17) 3 COMMON /AREA3/ WT(4,5,17,10) , SDET(4,5,17,10) , SSOMP (4,5,17,10) , S +PROD(4,5,17,10), SPRODP(4,5,17,10), SPROSP(4,5,17,10), SDETP(4,5,1 +7,10) 4 15 WRITE(6,1) 5 1 FORMAT(' ','ENTER: WTPAR(I1)1/'WET WT=1'/'DRY WT=2'/'AFDW=31) 6 RE AD (5, 2) WTPAR 7 2 FORMAT(II) 8 IF((WTPAR.GT.3) .OR.(WTPAR.EQ.0)) GO TO 15 9 16 WRITE (6,3) 10 3 FORMAT(' ','ENTER: PROB-LEVEL(F4.0) @ .01,.05 OR .10') 11 READ(5,4) PROB 12 4 FORMAT(F4.0) 13 X2=0. 14 IF(ABS(PROB-.01).LT..0001) X2=11.341 15 IF(ABS(PROB-.05).LT..0001) X2=7.815 16 IF(ABS(PROB-.10).LT..0001) X2=6.251 17 IF(X2.EQ.O.) GO TO 16 18 WRITE (6.14) PROB,X2 19 14 FORMAT('-','PROB LEVEL=',F4.2,3X,1X2=',F6.3) 20 CALL Ml 21 CALL M2 22 CALL M3 2 3 CALL M4 24 STOP 25 END 26 BLOCK DATA 2 7 COMMON /AREA1/ WTDAS(5,4,10) 28 DATA WTDAS/200*0./ 29 END 30 SUBROUTINE Ml 31 INTEGER SP,WTPAR,DAS2,DATE,DAS,TX 32 DIMENSION DAS1(4) 33 COMMON WTPAR /AREA1/ WTDAS(5,4,10) 34 DAS=2 35 DO 2 N=l,156 36 READ(2,4) DAS2,SP 37 4 FORMAT(7X,I3,6X,I2) 38 IF(N.EQ.l) DAS1(1)=DAS2 39 IF((SP.NE.3).AND.(SP.NE.6).AND.(SP.NE.7).AND.(SP.NE.8).AND.(SP.NE. +19) ) GO TO 2 40 BACKSPACE2 41 IF(WTPAR.EQ.l) READ(2,5) TXDX1,DATA 42 IF(WTPAR.EQ.2) READ(2,5) TXDX1,B1,DATA 43 IF(WTPAR.EQ.3) READ(2,5) TXDX1,B1,B2,DATA 44 5 FORMAT(10X,F3.0,6X,3F10.0) 45 IF(SP.EQ.3) SP=1 46 IF(SP.EQ.6) SP=2 47 IF(SP.EQ.7) SP=3 48 IF(SP.EQ.8) SP=4 49 IF(SP.EQ.19) SP=5 50 IF(DAS2.EQ.DAS1(DAS-1)) GO TO 6 - 163 -51 DAS1(DAS)=DAS2 52 DAS=DAS+1 5 3 6 DO 7 TX=1,10 54 7 IF((TXDX1.EQ.(TX*10.)-10.).AND.(SP.LE.5)) WTDAS(SP,DAS-1,TX)=DATA+ +WTDAS(SP,DAS-1,TX) 55 2 CONTINUE 56 RETURN 5 7 END 58 SUBROUTINE M2 59 COMMON WTPAR /AREA2/ WTDAS(5,4,10) 60 COMMON /AREA3/ WT(4,5,17,10), SDET(4,5,17,10), SSOMP(4,5,17,10), S +PROD(4,5,17,10), SPRODP(4,5,17,10), SPROSP(4,5,17,10), SDETP(4,5,1 +7,10) 61 INTEGER SP, TXDX1, TX, WTPAR, DATE, DAS 62 DIMENSION SUM1(5), SUM2(5), WT(5,10), FREQ(5,10), CHISQ(5,10), CHI +WT(5,4,10), STAND(5,10) 63 DATA SUMl/5*0./, SUM2/5*0./, WT/50*0./ 64 DO 1 N=l,625 65 IF(WTPAR.EQ.l) READ(1,2) TXTXl,SP,DATA 66 IF(WTPAR.EQ.2) READ(1,2) TXDX1,SP,B1,DATA 67 IF(WTPAR.EQ.3) READ(1,2) TXDX1,SP,B1,B2,DATA 68 2 FORMAT(10X,I3,3X,I2,1X,3F10.0) 69 IF((SP.NE.3).AND.(SP.NE.6).AND.(SP.NE.7).AND.(SP.NE.8).AND.(SP.NE. +19) ) GO TO 1 70 IF(SP.EQ.3) SP=1 71 IF(SP.EQ.6) SP=2 72 IF(SP.EQ.7) SP=3 73 IF(SP.EQ.8) SP=4 74 IF(SP.EQ.19) SP=5 75 TX=(TXDX1/10) 76 WT(SP,TX+1)=WT(SP,TX+1)+DATA 77 SUM1(SP)=SUM1(SP)+DATA 78 1 CONTINUE 79 DO 3 SP=1,5 80 DO 3 TX=1,10 81 FREQ(SP,TX)=WT(SP,TX)/SUM1(SP) 82 3 SUM2(SP)=SUM2(SP)+WTDAS(SP,1,TX) 83C ** CORRECTIVE ADJUSTMENT FOR AN UNREPRESENTATIVE DATUM FOR 'IRIDAEA 84C ** CORDATA' OBTAINED FOR THE 2 7 JULY 1976 COLLECTION AT 95 M. 85 SUM2(2)=SUM2(2)+20. 86 DO 4 SP=1,5 87 DO 4 TX=1,10 88 4 STAND(SP, TX) =FREQ (SP, TX) *SUM2 (SP) 89 DO 5 SP=1,5 90 DO 5 TX=1,10 91 UNIT=WTDAS(SP,1,TX) 92 CHIWT(SP,1,TX)=STAND(SP,TX) 9 3 DO 12 DAS=2,4 94 CHIWT(SP,DAS,TX)=WTDAS(SP,DAS,TX) 95 12 IF((WTDAS(SP,DAS,TX).LT.UNIT).AND.(WTDAS(SP,DAS,TX).NE.O.)) UNIT=W +TDAS(SP,DAS,TX) 96 IF(UNIT.EQ.O) GO TO 5 97 IF((WTPAR.EQ.l).AND.(UNIT.GT.10.)) UNIT=10. 98 IF((WTPAR.EQ.2).AND.(UNIT.GT.2.)) UNIT=2. 99 IF((WTPAR.EQ.3).AND.(UNIT.GT.l.)) UNIT=1. - 164 -100 SUM1=0. 101 SUMl=SUMl+(STAND(SP,TX)/UNIT) 102 IF(SUM1.EQ.0.) GO TO 5 103 DO 6 DAS=2,4 104 6 SUM1=SUM1+(WTDAS(SP,DAS,TX)/UNIT) 105 EXP=SUMl/4. 106 SUM2=0. 107 SUM2=SUM2+((STAND(SP,TX)/UNIT)**2) 108 DO 7 DAS=2,4 109 7 SUM2=SUM2+((WTDAS(SP,DAS,TX)/UNIT)**2) 110 CHISQ(SP,TX)=(SUM2/EXP)-SUM1 111 IF(CHISQ(SP,TX).GE.X2) GO TO 5 112 DO 8 DAS=1,4 113 8 CHIWT(SP,DAS,TX)=EXP*UNIT 114 5 CONTINUE 115 DO 11 DAS=1,4 116 DO 11 SP=1,5 117 WRITE (7,13) DAS, SP 118 13 FORMAT(1 -',3X,'DAS=',12,3X,'SP=',12) 119 DO 11 DATE=1,17 120 DO 10 TX=1,10 121 10 WT(DAS,SP,DATE,TX)=(REGWT(DATE)/REGWT(14))*CHIWT(SP,DAS,TX) 122 11 WRITE(7,9) DATE, (WT(DAS,SP,DATE,TX),TX=1,10) 12 3 9 FORMAT(' ',1DATE=',12,2X,10F10.4) 124 RETURN 125 END 126 SUBROUTINE M3 127 COMMON WTPAR /AREA2/ DAYl(17), P(ll), REGWT(17) 128 DIMENSION YRES(17), WT(17), SPRYY(411) 129 DIMENSION S(11), SIGMA(IO),'A(IO), B(10), DATEWT(17) 130 DOUBLE PRECISION YY(411), EXPO, RDATE 131 LOGICAL LK, ANSWER 132 INTEGER WTPAR,D,Y,DATE,DAS,TX 133 1 DATE=1 134 SUM1=0 135 REWIND1 136 DO 10 N=l,625 137 IF(WTPAR.EQ.l) READ(1,12) D,M,Y,DATA 138 IF(WTPAR.EQ.2) READ(1,12) D,M,Y,B1,DATA 139 IF(WTPAR.EQ.3) READ(1,12) D,M,Y,Bl,B2,DATA 140 12 FORMAT(312,13X3F10.0) 141 DAY2=JULDAY(M,D,Y+1900)-JULDAY(8,18,1975) 142 IF(N.EQ.l) DAY1(1)=DAY2 143 IF(DAY2.NE.DAY1(DATE)) GO TO 11 144 SUM1=SUM1+DATA/100. 145 IF(N.NE.625) GO TO 10 146 11 DATEWT(DATE)=SUM1 147 DATE=DATE+1 148 DAY1(DATE)=DAY2 149 IF(DATE.EQ.18) GO TO 10 150 SUM1=DATA 151 10 CONTINUE 152 NWT=0 153 K=10 154 N=17 155 LK=.TRUE. - ico -156 CALL OLQF(K.N.DAY1,DATEWT,REGWT,YRES,WT,NWT,S,SIGMA,A,B,SS,LK,P) 157 MAX=K+1 158 WRITE(7,2) (J, P(J), J=1,MAX) 159 2 FORMAT(' ',3('P(',12,')',E20.12,2X)) 160 WRITE (7, 3) K, SS 161 3 FORMAT(' ','K= ',I2,2X,'SS= ',F10.4/) 162 WRITE(7,4) (L, DATEWT(L), REGWT(L), YRES(L),L=1,N) 163 4 FORMAT(' ',2('DAY=' ,12 , ' DATEWT= ',F6.2,' REGWT= ',F6.2,' YRES= ', +F6.2,5X)) 164 WRITE (6,13) 165 13 FORMATC ','IS A PLOT OF *'TOTAL LITTER VS TIME'' DESIRED? (T OR F +) ') 166 READ(5,14) ANSWER 167 14 FORMAT(Ll) 168 IF(.NOT.ANSWER) GO TO 15 169 DO 5 DATE=1,411 170 YY(DATE)=0. 171 RDATE=DATE 172 DO 5 J=1,MAX 173 EXPO=J-l 174 5 YY(DATE)=YY(DATE)+(P(J)*(RDATE**EXPO)) 175 DO 9 DATE=1,411 176 9 SPRYY(DATE)=YY(DATE) 177 CALL SCALE(SPRYY,411,6.,YMIN,DY,1) 178 CALL AXIS(0.,0.,'1975*,-4,3.,0.,230.,40.) 179 CALL PLOT(3.,0.,3) 180 CALL PLOT(4. ,0.,2) 181 CALL AXIS(4.,0.,'1976',-4,7.,0.,25.,40.) 182 CALL AXIS(0.,0.,'LITTER BIOMASS (G/M2:AFDW)',26,6.,90.,YMIN,DY) 183 CALL PLOT(0.05,SPRYY(2),3) 184 DO 7 DATE=3,411 185 W=DATE*0.025 186 7 CALL PLOT(W,SPRYY(DATE),2) 187 DO 8 DATE=1,17 188 V=DAY1(DATE)*0.025 189 U= DATEWT(DATE)*0.0 2 190 8 CALL SYMBOL(V,U,0.28,30,0.,-1) 191 CALL SYMBOL(3.7,-.5,.2,'DAY OF THE YEAR',0.,15) 192 CALL SYMBOL(4.,5.,.2,'TOTAL LITTER' ,0 . ,12) 193 CALL PLOTND 194 15 RETURN 195 END 196 SUBROUTINE M4 197 COMMON /AREA2/ DAY1(17), P(ll), REGWT(17) 198 COMMON /AREA3/ WT(4,5,17,10), SDET(4,5,17,10), SSOMP(4,5,17,10), S +PROD(4,5,17,10), SPRODP(4,5,17,10), SPROSP(4,5,17,10), SDETP(4,5,1 +7,10) 199 INTEGER DATE1, DATE2, DATE3, DATE4, DATE5, DATE6, DATE7, DATE8, DA +TE9, DATE11, DATE12, SP, DAS, TX 200 DIMENSION DETP(523), SOMP(523), PROD(523), PRODP(523), PROSP(523), +DRATE(5), DET(523), DOM(5), YPROI(5), TEMFAC(523), SQLX(17) 201 DOUBLE PRECISION SUMl, SUM2, RATIO, PRATIO(ll), EXPO, DATE10 202 DOUBLE PRECISION QL, QLR(523), YWT(5,80), YWTI, YWTP, YWTC, YPRO(5 +,80), YPROP, YPROC, QLX(523) 203 DATA DRATE/.00760,.05651,.03123,.03479,.02934/, DOM/.393,.717,.589 +,.553,.611/, QLX/523*0./ - Ibb -204 DATA YPROI/.10892347,.12155714,.09405,.14905,.14466232/ 205 DO 13 DATE12=1,80 206 YWT(1,DATE12)=EXP((-.059039*DATE12)+4.60517) 207 YWT(2,DATE12) = (-.448099 *DATE12 **2)-(1,97802*DATE12)+100. 208 YWT(3,DATE12)=EXP((-.209873*DATE12)+4.60517) 209 YWT(4,DATE12)=((DATE12-6.022245)**2)/(4*.0906686) 210 YWT(5,DATE12)=EXP((-.277057*DATE12)+4.60517 211 YPR0(1,DATE12)=(-.067956*YWT(1,DATE12))+17.688 212 YPR0(2,DATE12)=(.0440286*YWT(2,DATE12))+7.75285 213 YPRO(3,DATE12)=(-.05 8322*YWT(3,DATE12))+15.2 371 214 YPR0(4,DATE12)=(-.182204*YWT(4,DATE12))+33.1254 215 YPRO (5,DATE12) = (.490 395E-0 3*YWT(5,DATE12)* *2)-(.21176*YWT(5,DATE12 +))+30.7386 216 13 CONTINUE 217 A=0.20187 218 B=0.29821 219 DO 8 DATE11=1,52 3 220 8 TEMFAC(DATE11)=1.375+A*SIN((8.*ATAN(1.)/366.)*(DATEll+231))+B*COS( +(8.*ATAN(l.)/366.)*(DATE11+231)) 221 DO 1 DAS=1,4 222 DO 1 SP=1,5 22 3 DO 1 TX=1,10 224 RATIO=WT(DAS,SP,1,TX)/(REGWT(1)*10.) 225 DO 5 1=1,11 226 EXPO=I 227 5 PRATIO(I)=(P(I)*RATIO)/EXPO 228 DO 12 DATE11=1,52 3 229 DETP(DATEll)=0. 230 SOMP(DATE11)=0. 231 PROD(DATE11)=0. 232 PRODP(DATE11)=0. 233 PROSP(DATE11)=0. 234 DET(DATEll)=0. 235 QLR(DATEll)=0. 2 36 12 CONTINUE 237 DO 16 DATE11=194,522 238 DATE1=DATE11 239 IF(DATEll.GT.410) DATEl=DATEll-407 240 DATE10=DATE1 241 SUM1=0. 242 SUM2=0. 243 DO 2 1=1,11 244 EXPO=I 245 SUMl=SUMl+(PRATIO(I)*(DATE10**EXPO)) 246 2 SUM2=SUM2+(PRATIO(I)*(DATE10+1.D0)**EXPO) 247 QL=SUM2-SUM1-QLR(DATE11) 248 IF(QL.LT.O-) QLX(DATE11+1)=QLX(DATE11+1)-QL 249 IF(QL.LE.O.) GO TO 16 250 IF(DATEll.LE.201) QLR(DATEll+l)=QL+QLR(DATEll) 251 YWPT=1.D0 252 YPROP=YPROI(SP) 253 DO 3 DATE2=1,80 254 DATE 3=DATE 2 +DATE11+(6 *TEMFAC(DATE11)) 255 IF(DATE3.GT.523) GO TO 16 256 DATE3=(TEMFAC(DATE3)*DATE2)+DATE11+(6*TEMFAC(DATEll)) - J.O / -257 YPR0C=YPR0(SP,DATE2)/100. 258 YWTC=YWT(SP,DATE2)/100. 259 IF((DATE3.GT.523).OR.(YWTC.LT..01)) GO TO 16 260 QLR(DATE3)=QL*YWTC+QLR(DATE3) 261 YWTI=YWTP-YWTC 262 IF(YWTC.GE.DOM(SP)) GO TO 4 263 DETP(DATE 3)=DETP(DATE 3)+(YWTI*QL) 264 PR0DP(DATE3)=PR0DP(DATE3)+YWTI*QL*((YPROC+YPROP)/2.)*(((YPROP*YWTP +)/YWTC-YPROC)/((YPROP*YWTP)/YWTC-YPROP 265 GO TO 10 266 4 SOMP(DATE3)=SOMP(DATE3)+(YWTI*QL) 267 PROSP(DATE3)=PROSP(DATE3)+YWTI*QL*((YPROC+YPROP)/2.)*(((YPROP*YWTP +)/YWTC-YPROC)/((YPROP*YWTP)/YWTC-YPROP 268 10 YPROP=YPROC 269 YWTP=YWTC 270 3 CONTINUE 271 16 CONTINUE 272 DO 15 DATE7=194,523 2 73 DO 11 DATE8=1,80 274 DATE9=DATE 7+DATE8-1 275 IF(DATE9.GT.523) GO TO 15 276 DATE9=TEMFAC(DATE9)*(DATE8-1))+DATE7 277 IF(DRATE(SP)*(DATE8-1) .GT.1.) .OR. (DATE9.GT.52 3)) GO TO 15 278 DET(DATE9)=DETP(DATE 7)*(1.-(DRATE(SP)*(DATE8-1)))+DET(DATE9) 279 11 PROD(DATE9)=PRODP(DATE7)*(1.-(DRATE(SP)*(DATE8-1)))+PROD(DATE9) 280 15 CONTINUE 281 DO 6 DATEll=412,52 3 282 QXL(DATE11-409)=QLX(DATE11) 283 DETP(DATE11-409)=DETP(DATE11) 284 DET(DATE11-409)=DET(DATE11) 285 PRODP(DATE11-409)=PRODP(DATE11) 286 PROD(DATE11-409)=PROD(DATE11) 287 SOMP(DATE11-409)=SOMP(DATE11) 288 6 PROSP(DATE11-409)=PROSP(DATE11) 289 SDET(DAS,SP,1,TX)=0. 290 SSOMP(DAS,SP,1,TX)=0. 291 SPRODP(DAS,SP,1,TX)=0. 292 SPROSP(DAS,SP,1,TX)=0. 293 SDETP(DAS,SP,1,TX)=0. 294 SPROD(DAS,SP,1,TX)=0. 295 SUM3=0. 296 SUM4=0. 297 SUM5=0. 298 SUM6=0. 299 SUM7=0. 300 DATE5=1. 301 DO 14 DATE4=2,411 302 SUM3=SOMP(DATE4)+SUM3 303 SUM4=PRODP(DATE4)+SUM4 304 SUM5=PROSP(DATE4)+SUM5 305 SUM6=DETP(DATE4)+SUM6 306 SUM7+QLX(DATE4)+SUM7 307 IF(ABS(DAY1(DATE5)-DATE4).GT..001) GO TO 14 308 S DET(DAS,SP,DATE5,TX)=DET(DATE 4) 309 SSOMP(DAS,SP,DATE5,TX)=SUM3 310 SPRODP(DAS,SP,DATE5,TX)=SUM4 - ibB -311 SPROSP(DAS,SP,DATE5,TX)=SUM5 312 SDETP(DAS,SP,DATE5,TX)=SUM6 313 SPROD(DAS, SP,DATE5,TX)=PROD(DATE4) 314 SQLX(DATE5)=SUM7 315 SUM3=0. 316 SUM4=0. 317 SUM5=0. 318 SUM6=0. 319 SUM7=0. 320 DATE5=DATE5+1 321 14 CONTINUE 322 1 CONTINUE 32 3 DO 7 DAS=1,4 324 DO 7 SP=1,5 325 DO 7 DATE6=1,17 326 WRITE(8,9) DAS,SP,DATE6,(SDET(DAS,SP,DATE6,TX),TX=1,10) 327 WRITE(10,9) DAS,SP,DATE6,(SSOMP(DAS,SP,DATE6,TX),TX=1,10) 328 WRITE(11,9) DAS,SP,DATE6,(SPRODP(DAS,SP,DATE6,TX),TX=1,10) 329 WRITE(12,9) DAS,SP,DATE6,(SPROSP(DAS,SP,DATE6,TX),TX=1,10) 330 WRITE(13,9) DAS,SP,DATE6,(SDETP(DAS,SP,DATE6,TX),TX=1,10) 331 7 WRITE(14,9) DAS,SP,DATE6,(SPROD(DAS,SP,DATE6,TX),TX=1,10) 332 9 FORMAT(II,11,12,IX,10E11.4) 33 3 DO 18 DATE6=1,17 334 18 WRITE(6,17) DATE6, SQLX(DATE6) 335 17 FORMAT(' ',12,2X,E11.4) 336 RETURN 337 END 

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