ABSTRAC Some characteristics of suspended particulate matter over two pocket sandy beaches were investigated. The amount of suspended material varied with wave turbulence but differences averaged out over time. Most particulate matter (80%) was algal detritus. Fifty per¬ cent ranged from .45 microns to 500 microns in diameter. A standing pool of littoral particuate matter seems to be maintained, alternating between a suspended and settled state. Availability to biota depends on balance between states. INTRODUCTION The sandy beach environment, though familiar to many, remains a complex and bewildering ecosystem. Its trophic structure is based on quantities of suspended particulate organic matter, which "may serve as food directly or by con¬ tributing to the total carbon budget" (Fox, 1950). The components of this suspended matter are phytoplank- ton, zooplankton, bacteria, and detritus. Numerous studes in various seas and coastal areas indicate that the major portion of particulate matter is detritus (Odum & de la Cruz, 1967, Krey, 1967, Khailov & Finenko, 1970, Finenko & Zaika, 1970, Suschenya & Finenko, 1966, Ryther & Menzel, 1965, Steele & Baird, 1961, Parsons & Strickland, 1962, Glynn, 1965). Detritus is here defined as "all types of biogenic material in various stages of microbial decomposition which represent a potential energy source for consumer species (Darnell, 1967), and is recognized as the energy source at the base of many ecosystems (Teal, 1962, Darnell, 1967, Heald, 1969, Odum & de la Cruz, 1963). Most of the organisms of the sandy beach (e.g. echino¬ derms, sipunculid & annelid worms) depend either on "copious supplies of finely divided and suspended, adsorbed, or preci¬ pitated organic detritus as a primary source of nutrition' (Fox, 1950) or on the larger algal detritus washed up as wrack. The concentration and composition of this organic matter at any time and place is subject to an intricate array of littoral factors, "an equilibrium between input rates and removal rates, and we can do little more than guess at these magnitudes" (Strickland, 1970). Though some work has been done on the role and composi¬ tion of organic material in estuarine and rocky intertidal environments (Glynn, 1965, McIntyre, 1970, Teal, 1962, Odum & de la Cruz, 1967, Darnell, 1967, Heald, 1969), very little research has been conducted on the energetics of the sandy beach ecosystem. This study is an attempt to determine the amount of suspended particulate matter (less than 1 cm in diameter) available to the sandy beach ecosystem in terms of organic carbon, the distribution of size and nature of the particles, the protein content of the particles, and some of the littoral factors affecting the availability of this material to the benthic community of the sandy beach. MATERIALS AND METHODS Two beaches at Hopkins Marine Station, characterized by vastly different physical parameters and ecological communi¬ ties, were chosen as sites for a comparative study. (Figure 1), Boatworks Beach is a long, shallowly sloping beach, dominated by fine sand grains (Md % - and protected from heavy wave action. Marinostat Beach is a shorter, steeply sloping beach composed of coarser sand grains (Md % -) and exposed to heavy wave action. Four two-liter sea water samples were taken at each beach just behind the roil line at approximately the same time, Preliminary studies corroborated those of Glynn (1965) indica¬ ting that there was no appreciable difference in samples taken during the light or dark periods of the day. Samples were therefore taken in daylight hours during the first 3 weeks of May at times subject to different tidal and other physical conditions. Local wave turbulence was estimated visually as the magnitude of water stirring at the roil line. The sea water was filtered through a series of Tyler screens with pores 1 cm, lmm, and 500 microns in diameter. A 500 milliliter aliquot of filtrate was passed through a .45 micron HA Millipore Filter coated with a suspension of 1% Mgcoz. The particulate matter remaining on the two smaller screens and millipore filter was washed into separate tubes and collected by centrifugation. Organic carbon was determined by a wet oxidation method (Strickland & Parsons, 1965). Protein was measured by the Lowry method. Chlorophyll a and phaeophytin were determined by flourometric analysis. (Strickland & Parsons, 1965). RESULTS Different quantities of suspended particulate organic matter were collected at Boatworks Beach and Marinostat Beach at different times. This variation was strongly correlated with the intensity of local wave stirring (Figure 2). Boat¬ works Beach was characterized by higher turbulence during the sampling time, resulting in slightly higher amounts of suspended carbon. Fluctuations in the abundance of sand were also related to water turbulence. When the amount of suspended material collected at each beach was averaged over the period of study, remarkably simi¬ lar figures were obtained: 2879 micrograms/liter + 1207 micro¬ grams/liter as one standard deviation (Boatworks Beach) and 2115 micrograms/liter + 1057 micrograms/liter (Marinostat Beach). This data is presented in Figure 3. The amount of suspended material in each of the three size classes considered: .45 microns - 500 microns; 500 microns¬ Imm; Imm - lcm; varied between the four sample sets taken at any one time on a beach and between beaches sampled at the same time. Thus, the relative proportion of material in each size class (and the corresponding standard deviation) was different each sample time (see data Appendix 1). Figure 4 gives some indication of this fluctuation in the size distribution of suspended organic carbon. The average amount of organic carbon in each size class was expressed as a percentage of the average total suspended organic carbon. Both beaches had similar size distributions (Figure 5). The smallest size fraction (.45-500 microns) con¬ tained the largest amount of organic carbon, the middle size raction (500 microns - lmm) contained the least. Microscopic examination of the suspended material pro¬ duced observations relatively consistent for the two beaches over time and corresponding closely with those of other studies (Glynn, 1965, Odum & de la Cruz, 1967). The pro¬ portions most commonly found were: Phytoplankton - 5%: Zooplankton - 7%; inorganic debris (e.g. sand) - 5%; and organic detritus - 80% (Figure 6). All but 2% or 3% of the organic detritus appeared to be fragmentary remains of benthic algal origin. The amounts of chlorophyll a (micrograms/liter) obtained for the smallest size class were converted to micrograms phytoplanktonic carbon per liter after the method of Finenko and Suschenya (1966). These values varied, both absolutely and as the percentage of total suspended organic carbon. A mean of 7% phytoplankton, correlating closely with values discussed above, was obtained for both beaches (Figure 6). Protein (percentage organic carbon) was determined for two different size classes: .45 microns to 500 microns, and 500 microns to 1 cm. The smaller sized material on both beaches contained much more protein than the larger particles, (Figure 7). DISCUSSION The composition and concentration of suspended particu¬ late organic matter in the water column over a sandy beach ecosystem vary constantly in time and space. These differences appear to be averaged out over a periodof time well within the lifespan of many sandy beach fauna. During the period of study, remarkable similarities were found in the amount and nature of material collected from two beaches characterized by different physical and biological parameters. This suggests that conditions other than the amount of suspended organic matter available to the ecosystem are important in determining the nature of some sandy beach communities. The size distribution of particulate organic carbon was virtually the same in both beaches. Approximately 53% of the material was in the size range .45 to 500 microns; 11% in the range of 500 microns to 1 mm; and 36% in the size class 1 mm to 1 cm in diameter. The predominant portion of suspended matter in both beach ecosystems was organic plant detritus (80%), followed by zooplankton (7%), phytoplankton (5%), and inorganic debris (5%). The smallest particles (.45-500 microns) were the richest in protein. A combination of several factors may be responsible: 1) the smaller organisms are protein-rich (e.g. phytoplankton, bacteria) and/or 2) the smaller detrital par- ticles are older and have a greater surface area to volume ratio, thus supporting a larger buildup of protein-rich micro¬ bial populations. Over the brief time period studied, it has been shown that Boatworks Beach and Marinostat Beach are exposed to essentially the same quality and quantity of suspended par¬ ticulate material. Plausible explanations include: 1) the source of input is the same and/or steady over time, 2) pro¬ cesses providing the material are similar, 3) the incoming material has been sufficiently mixed to disguise source variability. Both study sites are surrounded by the same rocky inter¬ tidal algal communities. These biologically rich areas could provide a substantial detrital contribution to the surrounding sandy beach ecosystems in the same manner reef detritus pro¬ vides a base for trophic systems within a lagoon (Marshall, 1965). The hydrodynamic forces which apparently cause extensive variation in suspended particulate organic matter at any one time and place are the same processes which would result in its steady input over time. They include surface winds, waves, currents, obstacles (e.g. rocks, larger algal drift), small benthic irregularities causing local eddies or pockets of calm, and the diminishing of littoral water forces with depth (Smith, 1968). During periods of calm, plant detritus usually settles with the fine-grained sediments. "But the topmost layer of such sediments is often flocculent and so easily stirred up that the detritus in it is continually re¬ exposed to decomposition in suspension" (Smith, 1968). Once the detritus is stirred, irregularities in water cells, the sheltering and filtering properties of floating wrack, and the ability of detrital particles to interfere with settling, all cause irregular pockets of detrital distribution along the beach (Smith, 1968). Greater amounts and variation in fine suspended particulate matter are thus to be expected under turbulent water conditions. This is clearly shown in Figure 2. On calmer days, when the littoral floor is free of stir¬ ring, "the final resting place of much of the plant detritus of the littoral is the littoral sediments" (Smith, 1968). There are many forces favoring the retention of this material within the littoral: the extent of the littoral, failure of water carrying properties with depth, and the settling ten¬ dencies of detritus, especially when further consolidated by mucus or transformation into fecal pellets. A picture emerges of a standing pool of littoral particulate material of rela¬ tively constant magnitude, alternating between a suspended and settled state. This pool is in equilibrium with overall littoral input and output rates. In the future, quantitative and qualitative studies should be conducted on this particulate littoral material, but its role in the energetics of the sandy beach ecosystem is of pri¬ mary importance and must not be overlooked: Basic information badly needed includes: 1) availability of material for feeding organisms. 2) actual utilization of the material under field conditions, and 3) nutritive value for the consuming species (Darnell, 1967). The precise nutritive role of organic detri¬ tus has yet to be determined. Though all the suspended material in the water column is theoretically "available" to consuming species, most of the benthic biota of sandy beaches can only utilize what is in, on, or close to the bottom. Of the standing pool of littoral particulate material, only a limited supply is abailable to any specific type of feeding organism aß one time. Critical tide factors divide the beach into sectors exposed to sudden sharp differences in water submersion time (Doty, 1971). Different lengths of exposure to water should divide the beach into sectors receiving distindly different amounts of nutrient material. The organic carbon content of sand from these different areas, and/or biotic communities characteristic of them, could be a function of critical tidal heights. The amount of organic matter entering and leaving the sandy beach ecosystemsis balanced against the amount retained within the system. As Seki (1968) observes, for a continuous input (L), where X - amount organic material remaining at timet X = (L/k) (1-e-kt decomposition rate factor (in sandy beach ecosystems, outflow and utilization rates are also components) If over long periods of time, X is a constant, steady state (ss) amount; Xss - L/k If one measures the organic carbon content remaining in the sand (Xgs) and L (the amt. suspended in the water) is the same for two pocket beaches, then k may be a useful index for comparing littoral sand beach communities. SUMMARY Some characteristics of suspended particulate matter, includ¬ ing amount, nature, and size distribution of the particles, as well as the littoral factors affecting its availability were investigated at two pocket sandy beaches in Monterey Bay. The amount of suspended material varied with wave turbulence, but differences averaged out over time. Most (80%) of the particulate matter was detritus of algal rigin. Zooplankton comprised 7% of the matter, followed by phytoplankton (5%) and inorganic debris (5%). The size distribution of particulate organic carbon was virtually the same in both beaches. Approximately 53% of the material was in the size range of .45 to 500 microns. The smallest particles were the richest in protein (70%). The hydrodynamic forces which apparently cause extensive variation in the organic material are the same processes which maintain a standing pool of littoral particulate matter. The alternation of this material between a suspended and settled state affects its availability to different sandy beach biota. ERENCES 1967. The organic detritus problem. Darnell, Rezneat M. IN Estuaries, G. Lauff (ed.), Am. Ass. Ad. Sci. Pub. No. 83:374-375. 1967. Organic detritus in relation to the estuarine ecosystem. IN Estuaries, G. Lauff (ed.), Am. Ass. Ad. Sci. Pub. No. 83:376-382 M.. 1971. Critical tide factors that are correlated Doty with the vertical distribution of marine algae and IN Readings other organisms along the Pacific coast. in Marine Ecology, J. Nybakken (ed.). Harper & How, New York:253-265. 1970. Particulate organic Finenko, Z. Z., and V. E. Zaika. matter and its role in the productivity of the sea. IN Marine Food Chains, J. H. Steele (ed.), Univ. of Calif. Press, Berkeley:32-14. Fox, D. L. 1950. Comparative metabolism of organic detritus by inshore animals. Ecology 31:100-108 Glynn, P. W. 1965. Community composition, structure, and interrelationships in the marine intertidal Endocladia muricata-Balanus glandula association in Monterey Bay, California. Beaufortia 118 (12). 1969. The production of organic detritus Heald, Eric J. in a south Florida estuary. Doctoral Dissertation at the University of Miami. Organic macro- 1970. Khailov, K. M., and Z. Z. Finenko. molecular compounds dissolved in sea water and their IN Marine Food Chains. inclusion into food chains. J. H. Steele (ed.):7-18. 1967. Detritus in the ocean and adjacent sea. Krey IN Estuaries, G. Lauff (ed.), Am. Ass. Ad. Sci. Pub. No. 83:389-394. Marshall, N. 1965. Detritus over the reef and its poten- tial contribution to adjacent waters of Eniwetok atoll. Ecology 16(3):343-344. 1970. Energy MoIntyre, A. D., Munro, A., and J. H. Steele. flow to a sand ecosystem. IN Marine Food Chains, J. H. Steele (ed.):19-31. Menzel, D. W., and J. H. Ryther. 1964. The composition of particulate organic matter in the western North Atlantic. Limnol. Oceanog. 9(2):179-186. Menzel, D. W., and H. H. Ryther. 1965. On the production, composition, and distribution of organic matter in the western Arabian Sea. Deep Sea Res. 12(2):199-209. Detritus as a major Odum, E., and A. de la Cruz. 1963. component of ecosystems. A.I.B.S. Bull. 13:39-40. 1967. Particulate organic detritus in a Georgia salt marsh-estuarine ecosystem. IN Estuaries, G. Lauff (ed.), Am. Ass. Ad. Sci. Pub. No. 83:383-388. Parsons, T. R., and T. D. Strickland. 1962. Oceanic detritus, Science 136:313-314 1965. A Manual of Sea Water Analysis. Fisheries Res. Board of Canada, Bull. 125. Parsons, T. R., Stephens, K. and J. D. H. Strickland. 1961. On the chemical composition of 11 species of marine phytoplankters. J. Fish. Res. Bd. Canada, 18(6). Seki, H. Skelding, J. and T. R. Parsons. 1968. Observations on the decomposition of a marine sediment. Limnol, Oceanog. 13:440-447. Smith, L. Kent. 1968. The role of benthic marine plants on the littoral phosphate cycle. Doctoral Dissertation at Stanford University, Steele, J. H., and I. E. Baird. 1961. Relations between primary production of chlorophyll and particulate carbon, Limnol. Oceanog. 6(1):68-78. Strickland, T. D. 1970. Recycling of Organic Matter. IN Marine Food Chains, J. H. Steele (ed):3-5. Suschenya, L. M., and Z. Z. Finenko. 1966. Concentration of suspended organic matter in tropical Atlantic water and some quantitative relationships of its components. Okeanologiya 6(5):835-847. Teal, J. M. 1962. Energy flow in the salt marsh ecosystem of Georgia. Ecology 13:614-624. ACKNOWLEDGEMENTS I would like to thank the faculty, staff, and students at Hopkins Marine Station for their help and encouragement. Special thanks go to Dr. Gilmartin, my advisor, to Dr. Phillips, for his (almost) unfailing good humor and aid, to Dr. Welton Lee for his brilliant bursts of intuition and helping hand, and to Delane Munson, without whose help this project would have been impossible. FIGURE 1 Beaches on which study sites were established, Mussel Point Pacific Grove, California. . Marinostat Beach C HOPKINS MARINE STATION meters Boatworks 90 Beach 0 7 JURE 2 Micrograms suspended organic carbon per liter versus wave turbulence at sample time. 5000 4000 53500 22500 o 2000 1500 1000 500 Marinostat Beach • Boatworks Beach — — FIGURE Average number micrograms suspended carbon per liter collected at each beach at different sampling times. and mean for each beach over entire sampling time. 0 2 2 S e ss . 1 — — c FIGURE 4 Amount organic carbon in each of 3 size classes at two study sites at two sample times. 100% 90 80 10 60 50 40 30 20 L Boatworks Beach Marinostat Beach I 5004- cm 454-500 —— 0 FIGURE! Size distribution of suspended particulate matter at study sites. 2400 2000 1600 1200 800 400 24 X 10" 20 12 b MAY 8 0815 MAY 8 1330 — Hüi m- 500u-mm C-45A-5004 . + W . — FIGURE 6 Composition particulate matter. Observed values are compared with those of Glynn (1965) and values calculated from chloro¬ phyll a tests. Bars mark range of values. C 5196 149 .. 5.. 1 35% . L Boatworks Beach 54% 89 : .. 38% . 9. Marinustat Beach — L . 454-500 5004-1 mm 1 mm-1cm 0 C FIGURE 7 Percent protein (of suspended organic carbon) for two size classes versus known values for other organisms. L — —. 8 S 5 —— —