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s Effects INTRODUCTION A vast amount of data and literature has accumulated s ince the 1975 NRC report. Much of this accumulation has been brought together at confer- ences (e.g ., Amer loan Petroleum Institute, 1975, 1977, 1979, 1981; Wolfe, 1977), at specialized symposia (American Institute of Biological Sciences, 1976, 1978; Fisheries Research Board, 1978; Cowan et al., 1981), and in two major reviews (Malins, 1977; Sprague et al. , 1981~ . This has come about in response to increased funding since the ear ly 1970s, partly due to increasing human concerns over oil in the marine environment, and partly as a natural outcome of continued spills and accidental discharges. One interesting and encouraging development has been a noticeable change in research emphasis, from descriptive to more process-or tented research, as in studies of physiological impact and ecological change (Table 5-1~. During the early days of oil pollution research following the Torrey Canyon accident, most research was aimed at quantifying toxicity thresholds. At the same time there was little scientific consistency, in that researchers developed their own exposure methodology and analytical preferences. AS a result, intercomparison of laboratory data was difficult. This began to change in the mid-1970s with a redirection of research interest toward understanding the mechanisms of hydrocarbon toxicity and the sites of toxic action. This effort was paralleled by concerted efforts of var. ious workers to standardize analytical methods, using certain reference oils set aside by the Amer ican Petroleum Institute (API). As a result, more meaning and comparability have come into the f ield of toxic e f feats of petroleum, and a type of data is being produced with which many members of the scientific community can agree upon (Rice et al., 19771. This change in emphasis in recent years represents a significant advancement since the 1975 NRC report. The f ield of oil pollution impact presents unusual and ma jor difficulties to the researcher in that at virtually every turn of study new techniques and analytical and sampling methods have to be devised. This is due to the newness of this research area, having come into its own only since 1967 with the breakup of the Torrey Canyon. Prior to that event most of the research interest with respect to petroleum concerned its physicochemical aspects, and the analytical methods arid 369

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370 TABLE 5-1 emphasis of Oil-Pollution-Related Study Reports for the Temperate and Nor thern Mar ine Environment Between 1967 and 1977 Bnphas is Pre-1974 Post-1974 1. Oil--physical-chemical changes, fate and distr ibution, env ironmental concentr ations 2 . Gross biological ef feats: mortality, toxicity Physiological, developmental, and ecological change Microbiology: hydrocarbon- utiliz ing banter ia 35% (83) 33% (107) 4396 (100) 31% (99) 6% (15) 2296 (70) 16% (37) 14% (44) NOTE: Numbers in parentheses denote number of studies. SOURCE: Environmental Protection Agency (1977 ~ . expertise reflected these interests. However, with the Torrey Canyon a new scientific discipline was required, including an understanding of the behavior of oil in water, sediment, and even in tissues, and requir- ing analytical methods capable of resolving petrogenic compounds in unfamiliar environmental samples . This has called for a much mor e interdisciplinary approach, with constant and new exchange of expertise __A .A~_~ Ion_ "~ ha: `:~_~. A.~.~1 ;~e. ~ ~ , ~ _, _ _ _ _ I -I I ~ ~- - - ~ - ~ - ~ -~~ - =- -..-- ~ Sc ientis ts in th is f ield are often competent in several areas, combining organic chemistry with biochemistry or biology, and often a more than passing acquaintance with microbiology or geology. Environmental teams of scientists have developed, working with integrated effort. This is not to say that the problem of understanding oil pollution in the mar ine environment is now well in hand . Ma jor inroads have been made in analytical capabilities and in understanding petroleum effects on two levels--physiological and ecological. However, generally there is a good appreciation of oil effects in temperate and northern temperate waters. At the same time the area of subcellular effects has received less attention. Ecological studies have been done pr imar fly in the field, using spills of opportunity and, more recently, large f ield enclosures (mesocosms) with known dosages of oil. At spills of opportunity, studies have been done mainly in the soft sediment areas such as salt marshes and shallow embayments. Such areas have been documented as sites where spilled oil will persist for long periods (years to

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371 decades) (see Chapter 3) and have provided the basis for most of what is known about how oil affects on mar ine populations and communities . The mesocosm studies have provided exper imental evidence of similar perturbations occurring in planktonic and benthic communities. Investigation into the physiological (e.g., photosynthesis, respiration, growth, neurotransmission) effects of oil is largely through laboratory investigations, although some fundamental work has been done in the field. One reason for the laboratory emphasis is the need to control experimental conditions. The coverage of this level of investigation has been uneven. AS in mammalian physiology, there are preferred invertebrate and algal species. Certain bivalves (Mya arenaria, Mytilus sp., Macoma spp.) and crustaceans (Cancer spp., Uca pugnax' Crampon spp., Penaeus aztecus) J by virtue of their accessibil- ity and ease of culture, are far easier to work with than benthic or pelagic organisms available only seasonally and/or by dredging or trawling. The same thing holds for the marine algae. It seems prefer- able, however, to attempt to understand in good detail the toxicology of petroleum in two or three well-studied representative organisms, rather than to attempt to establish simple toxic tolerance levels across all the phyla. (Although it should be noted that some of these may not be the most vulnerable.) Study at the subcellular level has received far less attention, despite recent concerns over certain hydrocarbons interacting with cellular macromolecules such as nucleic acids. We discuss primarily the impact of petroleum hydrocarbons, leaving the possible impact of other contaminants or compounds--either contained in oil or in some manner by-products of petroleum- or gas-related activities--to other more specialized discussions. Such contaminants would include, for example, trace metals, chemical dispersants, and drilling muds. Again, the available literature on petroleum impact alone is so vast that to include detailed discussions on these other materials, beyond merely mentioning them, would inevitably lead to an unmanageable exercise. The approach taken in this chapter is to discuss and review the impact of petroleum on marine biota and communities, by proceeding from one level ho the next--from effects on processes (cellular), through a discussion of effects on the marine foodchain (organismic), to the effects on communities (ecosystem). Inevitably this leads to some repetition or duplication, but this approach makes the most sense in unravelling and describing an extremely complex problem, involving a complex pollutant and the complexities of marine life. We hope that the index to this report will aid the reader in finding his or her way through it. Inevitably in a task such as this, some studies and reports will not have been referenced in the writing of this chapter because of the limited space available. Throughout we tried to refer to those studies that illustrated a particular point or aspect of petroleum pollution most aptly or most concisely. In other instances we referenced those studies that would lead the interested reader in turn to other studies. In an appendix to this chapter we have included a discussion of some well-known oil spills and oil seep problems, largely to add some

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372 real dimensions to the at times very detailed discussions in the main body of the chapter. The examples were selected for their general appropriateness and because they represented in each instance a particular spill type under certain conditions. Finally, any report tries to be up to date in its coverage, but inevitably there has to be a cut-off date. For various reasons, editor ial and technical, the review process for this chapter was very lengthy. We have tried to maintain as current a reference list as possible, but have not been able to go much beyond 1982-1983. We regret therefore having missed much excellent new literature and newer findings published in the last year. Toxicity In its most general sense, toxicity can be defined as the imparting of a deleterious effect, whether lethal or sublethal, to an organism, population, or community. The toxic effect can result in a permanent perturbation or change, for example, the crooked-back syndrome in larval fish, stunted growth, deformed shell formation in mollusks, and changed population patterns. However, not all effects are disruptive, and there exist adaptive mechanisms, both at the cellular level (e.g., detoxifying enzyme systems) and at the population and community levels (Capuzzo, 1981~. Toxic effects from petroleum exposure vary widely and for reasons that are not well understood. Certainly these have to do with the complexity of its chemical composition, with different products or even crude oils differing markedly in their chemical makeup. Another factor is the var lability in sensitivity to oil found among mar ine organisms, differing not only with the species (Figure 5-1) but even for life- cycle stages (Figure 5-21. While it is generally true that the younger stages of organisms are more sensitive to petroleum hydrocarbons, there are exceptions. Unfortunately, not many studies have compared the sensitivity of organisms at various life stages under identical experi- mental conditions for any one species. While the absolute toxicity of petroleum hydrocarbons appears to be greater for the higher-molecular-weight compounds (for example, 3- and 4-ring aromatics), most of the toxic effect of petroleum in water is thought to be due to the lower-molecular-weight (C12-C24) e-paraffin compounds and to the monoaromatic fraction, for the simple reason that these compounds are the most water-soluble (Chapter 3, Table 3-4~. From examinations of the concentrations present in water-soluble fractions (WSF), it is clear that the contribution of compounds higher in molecular weight than the alkyloaphthalenes is very small and may be insignif icant in terms of producing acute toxicity. Bioassay tests have been used to a considerable extent to determine the toxicities of var ious crude oils and of refined products. Most of these tests have used mortality as the index of toxicity, expressed for example, as LC5g (the lethal concentration yielding 50% mortality over pre-determ~ned exposure time, for example, 24, 48, or 96 hours). In practice, however, their usefulness as a research technique is

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PR] 26 24 22 20 18 ~ 16 IS 14 t`' 1 2 8 4 3 3 373 3 n I Crago 2 Neanthes 3 Cancer 4 Salmon fry 5 Cyprinodon 6 Copepod 7 Palaemonetes 8 Amphipod 9 Striped bass 10 Penaeus aztecus 11 Grass shrimp 24 hour LC50 5 ~ ~ -I - - ~ Jo ; ~ '' '''aft " ~1~111151 ' Ill~llllt~ly ~J ~ ~ 7~ _~ n ,~e, ~ 0~ ~ ~ ~ `~ AQUA O`~ Opt O`= 6~ O'er ~ O~ 10 ~ ~ it_ ~~s Cal FIGURE5-1 Acute toxicity (24- and 96-hour LC50 static tests ~ of some aromatic hydrocarbons for selected mar ine macroinver tebrates and fish. SOURCES: Caldwell et al. (1977), Benville and Korn (1977), Neff et al. (1976), R.E. Thomas and Rice (1979), Young (1977), Rossi and Neff (1978), Ott et al. (1978), and W.Y. Lee and Nicol (1978).

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374 A; ~ I: , ~ ~ ~ At. FIGURE S-2 Effects of oiling on f ine structure of surf smelt embryo retinas. (Left) Retinal cells of an unoiled embryo (x5000~. Inset is enlargement of a synaptic junction X18,000. {Right) Retina' cells of an embryo exposed to 113 ppb Cook Inlet crude oil (xSOOO). vesiculation is evident in the myoid regions (asterisk) of the receptor cells. Note also necrotic neurons (arrows). Synaptic junction (viz., inset, x18 '000) appears normal. Ellipsoid region (e) of inner segment of receptor cell' nucleus {m) of receptor cell' outer segment (o) of receptor cell, and synaptic (s) junctional complex are indicated on figure. (Photo by J. Hawkes.) limited in that they provide no data except on mortality. Toxicity tests are subject to several variables such as complex mixture of the oil, test parameters, and various biological factors such as age, sex, and contamination history of the organism. For these reasons they are somewhat imprecise measures of toxicity, and many researchers feel that their output, the LC50, has little relevance to what may happen to an organism as a result of a spill. By its very nature the LC50 gives no indication of sublethal toxic problems that the organism may be experiencing, and gives no measure of any long term impacts that may be occurring, measuring only the death of the organism. Instead, acute lethal bioassays serve best as tests to compare the relative toxicities of complex, unknown toxicants, or the comparison of relative sensitivities of species or life stages.

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375 Laboratory Versus Field Studies Most studies of effects have been developed in the laboratory for the simple reason that field studies often depend on spills of opportunity, which are highly unpredictable . ALSO, f ield studies are often expen- sive and difficult to carry out. On the other hand, laboratory studies have frequently been criticized because experimental conditions do not simulate field conditions. Also, concentrations of oil or hydrocarbons frequently exceed those encountered in the field. Both criticisms are probably correct. However, in recent years attempts have been made to bring laboratory conditions closer to field conditions by simulating the hydrocarbon composition and concentrations more closely through the use of flow-through systems and by careful management of the test organisms chosen. Most promising in this respect have been various studies carried out in "mesocosms," large enclosures that allow control of some environmental variables under near-open ocean conditions (for example, Marine Ecosystems Research Laboratory (MERL), CEPEX, Loch Ewe, viz., Table 5-~. In this respect, natural oil seeps also offer certain opportunities to study the impact of petroleum under more open-ocean conditions. As for the criticism of high experimental dosages, there are situations where seemingly high concentrations of oil or hydrocarbon are warranted, for example, in the initial establishment or detection of certain toxic effects and in the analysis of metabolic pathways. High initial concentrations of toxicants are frequently necessary to establish a toxic effect that otherwise might be indistinguishable from the background "noises or masked by other changes. Using high concen- trations allows the experimenter to better define the toxic effect or response. Again, the detection and identification of primary or secondary metabolites, or of short-lived intermediates, often requir e unusually large doses of toxicant. In these instances the object is not so much to determine a toxic effect, as it is to better understand certain aspects of hydrocarbon metabolism for which low dosages would not elicit a measurable response. However, care has to be taken in , work ing with h igh dosages and in interpreting results because of the possibility that extraordinary metabolic pathways may be expressed. In the end, both laboratory and field study have merit. Although field studies are fraught with uncontrolled and interfering factors, they nonetheless are the ultimate testing ground. For that reason, spill sites should be visited, and revisited, whenever possible. On the other hand, laboratory studies support field studies by providing the opportunity to investigate an effect in detail and to study its under- lying mechanism. Factors Affecting Impact of Oil When an oil spill occurs, many factors determine whether that spill will cause heavy, long lasting biological damage; comparatively little or no damage; or some intermediate degree of damage. An example of the variability that exists among the effects of oil spills on the mar ine

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376 biota is outlined by C.T. Mitchell et al. (1970) in their description of the widely different effects resulting from the Tampico Maru and the Santa Barbara oil spills. Nonetheless, there are some patterns begin- ning to emerge that are useful in identifying those physical and biological features that can influence the ultimate impact of a spill or chronic pollution (e.g. , Michael et al. , 1978; Gundlach and Hayes, 1978; Owens and Robilliard, 1981; Vandermeulen, 1977, 1982) . Geographic Location In many reports, organisms from any one geographic location are apparently no different from any other location in terms of their vulnerability to petroleum hydrocarbons. While there are of course species-specific genetic differences, arctic fish or invertebrates do not appear to differ physiologically from similar organisms at lower or tropical latitudes in terms of lethal toxic concentration thresholds or toxic r esponses . However, there are a number of physical features related to geo- graphic location--mainly temperature and ice cover, together with differences in community diversity, which is latitude dependent--that will influence both impact and biolog i Cal recovery. Temperature, for example, plays a significant role in the solubility of hydrocarbons in the water column and in the rate of their degradation through microbial activity. Similarly, community diversity at different latitudes (low diversity in polar regions, high diversity in tropical environments) can lead to differences in both times and patterns of biological recovery following an oil impact. Oil Dosage and Impact Area If the spill occurs in a small, confined area so that the oil is unable to escape, damage will be greater, almost without exception, for a given volume and type of oil spilled than if that same volume were released in a relatively open area. For example, at the Arrow spill site in Chedabucto Bay, Nova Scotia, about 2.5 million gallons of Bunker C fuel were spilled in an embayment, whereas the Argo Merchant spilled about 7.7 million gallons of No. 6 fuel oil into the open ocean of f Nantucket Island, Massachusetts. Although a considerable amount of Arrow spill eventually was swept out to sea, the confined nature of the oil during the first days resulted in nearly uniform oiling of the entire bay coastline and in considerable damage to the associated fauna and flora. However, this generalization is not inflexible. The spill of the super- tanker Amoco Cadiz occurred offshore, but prevailing winds were such that the oil was kept near shore of North Brittany for several weeks and continuously driven onto shore. Similarly, there are differences resulting from the manner of the spillage--whether low level but chronic or consisting of a sudden accidental release. The former is covered in greater detail elsewhere (for example, see Chronic Oiling section), but

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377 generally the impact from such chronic releases differs both in sever ity and in k ind from accidental spills, where the spilled oil will eventu- ally disappear with time due to physical-chemical processes and micro- b ial and other biological degradation . On the other hand, in the case of chronic releases the spilled oil becomes a continuing irr itant or toxicant to which the community must adjust, for example, in selection for hydrocarbon-utiliz ing species . Oceanograph ic Conditions Currents, sea state, coastal topography, and tidal action all combine to influence the impact of a given spill. Currents and wave action--in open water or open bays--act to break up the oil into smaller slicks, and also act to disperse some of the oil into the water column. In areas of large tidal ranges the oil can become distr ibuted over a broad range of the intertidal zone, arid can be deposited far above the high tide mark by extreme "spr ing" tides coinciding with high winds and strong tidal flow . Coastal topography plays a large role in the residual impact of a spill, with low energy environments (salt marshes, lagoons, estuaries, embayments) acting as long term hydrocarbon "sinks." Impact on biota in such systems is usually long lasting. Meteorological Conditions Normally, storms increase wave action and wind speed and thereby aid in evaporation of the lower-molecular-weight, more volatile toxic compo- nents . On occasion, however, wave action may intensify the problems, as apparently occurred at ache Flor Ida No. 2 fuel oil spill near West Falmouth. Soon after this spill, the surf drove the oil ashore into the sediments and the surrounding marshland (Sanders, 1978) . The oiled marshland and sediments then became a long term reservoir of oil with persistence in some areas to this day. A1SO storm-induced resuspension of subtidal sediments probably brought these sediments into contact with oil from more intertidal areas. Similar events occurred following the breakup of the Amoco Cadiz, where the winter storms drove the oil deep inland up the nearby estuar ine tidal r ivers (Hess, 1978 ~ . Season Season is particularly important in terms of the biota that might lie in the path of a spill or in the vicinity of a chronic oiling s~tua- tion. For example, if a spill occurs in an area where seabirds are feeding or nesting, bird mortality might be in the thousands; at some other time of the year the mortality might be much lower. Similarly the coincidence of a spill with f ish spawning events or hatching and development of larval f ish migration might result in higher than normal larval mar tall ties .

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378 Seasonal changes in the phys decal parameter s of the mar ine environ- ment might also influence the potential impact of a spill, as for example, seasonally timed changes in local circulation patterns that might lead to local containment of slicks. Oil Type Oil type determines both the short term and the long term impact. Immediate impact can be very high from such highly toxic oils as diesel and jet fuel. However, these dissipate readily and leave relatively little residue, unlike the crude and Bunker oils, which can persist in certain sediments for up to several decades. This aspect is not as simple and clear cut as it seems, however. Traces of No. 2 fuel oil, a relatively volatile product, still persist in sediments of Falmouth, Massachusetts, 13 years after the spill of the Flor Ida . Oil Metabolites and Photochemical Reaction Products This subject deserves separate mention as it was not raised in any detail in the 1975 NRC report, but has become of interest in more recent years, following observations that some petroleum metabolites or intermediate products can be quite toxic and may even have mutagenic proper ties . Oil metabolites can be formed from the parent oil by biological conversion of compounds taken up by marine biota (including bacteria), and by photochemical processes. The relatively few data available on either method come mainly from laboratory studies. The formation of compounds by photochemical processes has been addressed earlier (Chapter 4 ~ . In general, irradiated samples of petroleum or water-soluble preparations appear to be more toxic than the parent compounds. For example, Lacaze and Villedon de Naide (1976) cite field studies sug- gesting that the irradiated water-soluble fractions (WSF) of Kuwait crude oil were 3 times as toxic, depressing C-fixation, as nonirradiated WSF after 40- and 64-hour exposure of the alga Phaeodactylum cornutum. In similar studies, Scheier and Gominger (1976 ~ examined the toxic effects of irradiated versus nonirradiated No. 2 fuel oil, using a Sylvania sunlamp, and compared the results with solar-irradiated WSF. They observed that (1) sunlight was nearly 10 times more effective than sunlamp exposure in raising the toxicity of the irradiated WSF, as indicated by the anthracene-dianthracene conversion ratio, and (2) both significantly increased the toxicity of the WSF due to the irradiation. There is 1 ittle or no information on the potential toxicity of bio, ogical metabolites of petroleum compounds, and any conclusion is difficult, for metabolites have been demonstrated in only a few instances (e .g ., Corner and Harr is , 1976 ; Sanborn and Mal ins , 1977, 1980; Varanasi and Gmur, 19801. There is no evidence to date that the bulk of the petroleum hydrocarbon metabolites formed by biological activity are any more toxic than their parent compounds e However, a small proportion of petroleum compounds do give rise to mutagenic

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379 intermediates and to metabolites capable of binding with nucleic acids (Varanas~ and Gmur, 1980; Varanasi et al., 1980, 1982~. This potential appears to be limited only to the polycyclic aromatic hydrocarbons. The evidence to date is sparse but does not indicate that a mutation load has been introduced generally into the mar ine environment by this mechanism as a direct result of petroleum spillage or chronic spillage. However, this possibility cannot be ruled out in isolated incidents. Remedial Measures A great deal of effort continues to be expended, on countermeasures and various cleanup and control methods. These generally fall into one of two categories--mechanical and chemical--and because of their nature they inevitably leave some traces on the landscape, be it some form of physical disruption following mechanical cleanup or the risk of chemical alteration following application of chemical methods. As these are an almost automatic response to oiling incidents, a brief discussion of their potential effects on the marine environment seems appropriate. Mechanical Containment and Cleanup This category includes those methods which focus on the actual removal of oil or oiled debris, as by bulldozing or hosing with water under pressure. Most of this activity involves the intertidal zone. Offshore oiling incidents rarely are suitable for mechanical cleanup except by surface skimmers or possibly the cropping of oiled kelp using mechanical aquatic weed cutters. Neither of these is very likely to have much of an adverse effect on the environment. However, the problem becomes more serious in the intertidal zone, largely due to the physical disrup- tion of habitats. Rocky coastlines present the least problem in terms of cleanup. Oiled rocky surfaces are cleaned most often with either flushing, steam cleaning, sand blasting, or manual scraping. None of these is likely to alter the substrate to any extent, and the main damage is the removal of fauna and flora. The biological recovery process may take several years, but nevertheless, recovery will occur. As the settling surfaces have probably not been chemically or physically altered in the cleanup process to any great extent, the only limitations to recovery are biological ones. Of course, the rerelease of the stranded and flushed oil into the water column may pose additional problems. The problem becomes greater with the oiling of finer-grained sediments such as cobble-boulder beaches or the fine silt sediments of lagoons and marshes. Because of the penetration of oil into such sediments, removal of oiled sediments often accompanies cleanup. Excessive removal can result in the disturbance of physical and ecological equilibrium. Excessive removal of beach sediments can lead to beach retreat or beck shore (cliff) erosion. This was observed following the Arrow disaster, where a 20-m landward movement of pebble- cobble beach was recorded following large scale removal of oiled cobble

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538 Smith, T.G., and I. Stirling. 1975. The breeding habitat of the r inged seal (Phoca hispida) . The birth lair and associated structure. Can. J. Zool. 53 :1297-1305. Smithsonian Institute. 1970a. S. I. Event No. 24-70. Center for Short-lived Phenomena, Washington, D.C. Smithsonian Institute 1970b. S. I. Event No. 186-70. Center for Short-lived Phenomena, Washington, D.C. Smithsonian Institute. 1972. S. I. Event No. 1-72. Center for Short-lived Phenomena, Washington, D.C. Snedaker , S. C., J.A. Jimenez , and M. S. Brown. 1981. Anomalous aer ial roots in Avicennia germinans (L.) in Florida and Costa Rica. Bull. Mar. Sci. 31~2) :467-470. Snow, N.B., and B.F. Scott. 1975. The effect and fate of crude oil spilt on two arctic lakes, pp. 527-534. In Proceedings, Joint Conference on Prevention and Control of Oil Spills. Amer loan Petroleum Institute, Washington, D.C. Soto, C., J.A. Hellebust, and T.C. Hutchinson 1975a. Effect of naphthalene and aqueous crude oil extracts on the green flagellate Chlamydomonas angulosa. II. Phytosynthesis and the uptake and release of naphthalene. Can. J. Bot. 53:118-126. Soto, C., Hellebust, J.A., Hutchinson, T.C. and Sawa, T. 1975b. Effect of napht~halene and aqueous crude oil extracts on the green f lagellate Chlamydomonas angulosa. I . Growth. Can. J. Bot. 53:109-117. Southward, A.J. 1982. An ecologist's view of the implications of the observed physiological and biochemical effects of petroleum compounds on mar ine organisms and ecosystems. Phil. Trans. R. Soc. London, Ser . B 297: 241-255 . Southward, A.J., and E.C. Southward. 1978. Recolonization of rocky shores in Cornwall after use of toxic dispersants to clean up the Torrey Canyon spill. J. Fish. Res. Board Can. 35~5) :682-706. Sousa, G. 1970. Report of the Shellfish Warden, pp. 161-165. In Annual Repor t of the Finances of the Town of Falmouth for the year ending December 31, 1970. Spaulding, M. L. 1978 . Sur face and subsur face spill tra jectory forecasting: application to the Argo Merchant, pp. 37-42. In M. P . Wilson, J. G. Quinn, and K. Sherman, eds . In the Wake of the Argo Merchant. Center for Ocean Management Studies, University of Rhode Island, Kingston. Spies, R.B., and P.H. Davis. 1979. The infaunal benthos of a natural oil seep in the Santa Barbara Channel. Mar. Biol. 50 :227-238 . Spies, R.B., and P.H. Davis. 1981. Toxicity of Santa Barbara seep oil to start ish embryos. III . Influence of parental exposure and the effects of other crude oils. Mar. Environ. Res. In press. Spies, R.B., and P.~. Davis. 1982. Toxicity of Santa Barbara seep oil to starfish embryos. 3. Influence of parental exposure and the effects of other crude oils. Mar . Environ. Res . 6: 3-11. Spies, R.B., and D.J. DesMz~rais . 1983. A natural isotope study of trophic enrichment of marine benthic communities by petroleum seepage . Mar . Biol . ~ in press ~ .

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