sign in sign up
<<
Home , Barnes Sound, Captiva Pass, John Hart (actor), Predatory dinoflagellate, Red tide

535^

Red- Research Summarized to 1964 Including an Annotated Bibliography

By George A. Rounsefell and Walter R. Nelson

Marine Biological Laboratory LIBRARY MAR 2 2 1967

WOODS HOLE, MASS.

SPECIAL SCIENTIFIC REPORT- Na 535

UNITED STATES DEPARTMENT OF THE INTERIOR Stewart L. Udall, Secretary Charles F. Luce, Under Secretary Stanley A. Cain, Assistant Secretary for Fish and Wildlife and Parks

FISH AND WILDLIFE SERVICE, Clarence F. Pautzke, Commissioner

Bureau of Commercial Fisheries, Harold E. Crowther, Acting Director

Red-Tide Research Summarized to 1964 Including an Annotated Bibliography

By

GEORGE A. ROUNSEFELL and WALTER R. NELSON

Contribution No. 20, Bureau of Comnnercial Fisheries Biological Laboratory, St. Petersburg Beach, Fla.

United States Fish and Wildlife Service Special Scientific Report--Fisheries No. 535

,4 Washington, D.C. Decennber 1966

CONTENTS

Page

Introduction 1 Historical 2 General conditions during red-tide outbreaks 3 Temperature 3 3 Rainfall 4 Wind 4 Light 4 5 Nutrient levels 5 Are preceding blooms conducive to red tide? 5 Distribution of red-tide bloonns 6 Effect of dead fishes in continuing a red-tide bloom 7 Field observations on red tide 7 Seasonal occurrence of red tide 8 The role of convergences in causing red-tide blooms 8 Coastwise and seasonal abundance 10 Pollution as a contribution to red-tide outbreaks 10 Life forms of breve 10 Control of red tide 12 Damage from red-tide outbreaks 12 Methods of control • • 13 Prevention of outbreaks 13 Control of outbreaks 13 Suggestions for future research 14 Laboratory studies of the organism 15 Field studies of red tide 16 Control methods 16 Need for coordinated research 16 References pertaining to red tide and related subjects 17 Subject index to selected topics 76 Appendix table 1 85 Frontispiece. —A representative sample of dead fish deposited on beach in the lower part of during the 1963 red tide. Red-Tide Research Summarized to 1964 Including an Annotated Bibliography

By

GEORGE A. ROUNSEFELL and WALTER R. NELSON

ABSTRACT

This paper summarizes from published and unpublished data and reports the status of research on the Florida red tide up to 1964. It contains 292 references, mostly annotated, on red tide and closely related subjects. The relation of oceano- graphic conditions to red-tide blooms, the seasonal and coastwise distribution of the Florida red tide, and progress in various aspects of research are discussed.

INTRODUCTION allotted time. The list includes many refer- ences which, although not examined by us, This summary of red-tide research is a may prove useful to anyone needing closely modified version of a report prepared for the related infornnation. Stress was placed on Bureau of Commercial Fisheries under Con- material relating to the red tide of Florida. tract No. 14-17-0002-79 with the University No attempt was made to undertake a synopsis of Alabama {Status of red-tide research in of each paper; rather the comments and ex- A. Rounsefell and Walter R. 1964, by George cerpts given represent a deliberate attempt to 64-1, Alabama Ma- Nelson, Technical Report bring out important facets of the research Laboratory, 192 rine Resources 1964, p., problem. Over 20 new references have been processed). added to the 1964 report. The aim of the first report was to examine As a second step, the senior author is col- and review published and unpublished papers laborating with staff mennbers of the Bureau and data relating to red tide, to summarize of Commercial Fisheries Biological Labora- this material, and to make the summary tory in St. Petersburg Beach, Fla., in prepar- available prior to a symposium on red tide ing a paper based on 7 years of published raw called by the Bureau in October 1964. A data on red tide covering the period 1954-61 similar symposium, held at the Bureau of collected by the staff of the Laboratory. The Commercial Fisheries Biological Laboratory data were coded, punched, and verified on in Galveston, Tex., March 5 to 7, 1958^, re- nearly 13,000 ADP cards and run through the sulted in several modifications to improve the solid-state Univac-80 computer at the Univer- then existing research program. We hope that sity of Alabama Computer Center in Tusca- the 1964 symposium has also been useful in loosa in preliminary analyses, some of which assessing current progress and suggesting are presented in this report (see fig. 1). promising modifications of the research pro- Finally, we presented our ideas on the gram. rationale of red-tide occurrence, together began by reviewing as of the We many with suggestions for future research. This published papers and manuscripts on red tide section of our report was presented with as could be obtained and examined in the some trepidation in view of the expert audience Note.— George A. Rounsefell, Professor of Marine at the symposium. It represented ideas gained Biology, and Walter R. Nelson, Marine Biologist, Uni- chiefly from winnowing of the extant reports, versity of Alabama, University, Alabama. 35486 and we hoped that the participants in the •'•All excerpts or references from the report of this symposium would not regard it as an attempt earlier symposium (mimeographed) are identified simply to push our ideas, but as an attempt to furnish as (USFWS, 1958). material for stimulating discussion. Figure 1 .--North-South areas used In coding Florida red- tide data.

HISTORICAL the tremendous bloonns of the diatom Axila- codiscus kittoni at Copalis Beach, Wash. A Red tide with its concomitant mass nnortali- few discolorations arise from swarms of ties is not a phenomenon peculiar to Florida. , from , or from chromogenic Overblooming of organisms has been bacteria, but the great majority are caused well known since ancient times. It should by , some of which are appar- suffice to note that discolored water has been ently only slightly toxic, if at all. They vary observed in widely scattered ; the Red in color from almost transparent, through and the Gulf of (once nanned pale green and yellow, to amber and bright the Vermilion Sea) are cases in point. Many red. Many are luminescent; some have a discolorations are apparently harmless, or at definite cuticle (are armored), and many are least caused by nontoxic organisms, such as naked. In some localities the overblooming occurs only sporadically and reports are few. remain fairly low in regions of extensive up- The senior author found no reports, for in- welling, such as the California and southwest stance, from Alaska, although he knows from African coasts, where red often occur. personal experience that streaks of water as Barker (1935) found the optimal range for 14 red as tonnato soup sometimes occur in species of marine dinoflagellates to lie between southeastern Alaska in late summer. 18° and 35 C. For Gymnodinium breve Finu- The best docunnented reports of red tides cane (1960) stated that the optimum is 26° to and mass mortalities conne from a few locali- 28° C, but that dense populations were ob- ties: The Malabar Coast of India, the coast served at 15° to 18° C. Aldrich (1959) exposed of northern Chile and Peru, near Walvis Bay 800 individual cultures to controlled tempera- in Southwest Africa, Japan, and southern tures and found no survival at 7° C. or below, California. Additionally, in many localities in or at 32° C. and above. Populations did not which blooms of toxic dinoflagellates are not grow above 30° C, and multiplication was sufficiently dense to cause mortalities, the visibly slowed but not halted at 15° C. ingested organisnas can render toxic Isopleths of abundance of G. breve against to humans. tennperature and salinity (averages from The distribution of the naked , samples of 20 or nnore) from 7 years of field

Gymnodinium breve , that causes the Florida data analyzed on the computer show that the red tide is not well known. It occurs in organisms appear to thrive from 16 to 27 C. Trinidad (Lackey, 1956) and is the chief (fig. 2). Beyond this range organisms occurred suspect in frequent fish mortalities near the only in low abundance. These observations mouth of the Orinoco River in the Gulf of agree rennarkably well with the laboratory Paria. There is considerable evidence that results of Aldrich (1959). G. breve may be distributed throughout the The effect of cold waves in ending a bloom Gulf of . The organism has caused has often been mentioned. After the heavy out- extensive fish nnortalities along the southern break in the fall of 1957, Finucane (USFWS, coast of Texas, extending far down the Mexican 1958) said, "...There were several cold waves coast (Wilson and Ray, 1956; Gunter, 1952; in December and water temperature was as Lund, 1935). Wilson isolated G. breve from low as 9.9° Centigrade. Following the cold front Galveston waters (Collier, 1958b). Fishermen we couldn't find any bugs north of Pass-a- report "red water" on the Campeche Banks, grille. . . ."Taylor ( 1917a, b) mentioned a severe but the causative agent is still unknown freeze in February 1917, following the heavy (Graham, 1954). Because G. breve was not fall outbreak of 1916, which did not recur in identified until 1948 and is so fragile as to 1917. Gunter (USFWS, 1958) mentioned a resist collection by conventional methods, it cold spell that brought freezing temperatures may be far more widely distributed than avail- to Miami and ended (temporarily at least) able records indicate. Although other toxic the 1946 red tide outbreak.

dinoflagellates, such as , occur in Florida, it is probably safe to assume that Salinity the earlier red tides were also caused by Dinoflagellates in general have a wide salin- C_. breve . Hutton (1956) reported a bloom of ity tolerance. The armored species are usually furca with counts as high as 17,600,000 cells per liter, but this was in Hillsborough Bay (a part of Tampa Bay) which is not normally invaded by G. breve on account of the low salinity (see Ingle and Sykes, 1964),

GENERAL CONDITIONS DURING RED-TIDE OUTBREAKS

Conditions conducive to overblooming of dinoflagellates are suggested by numerous reports of red-tide outbreaks in many locali- •ties. These are discussed below.

Temperature

Red tides as a whole are less frequent and less intense in temperate regions than in the lower latitudes, where the hot weather during outbreaks is often mentioned. These observa- tions concerning hot weather may be a reflec- Figure 2.— Isopleth (0.01) of geometric mean of Gymno- tion of other conditions, such as a lack of dinium breve per ml. showing relation of G. breve to strong wind. Certainly the sea temperatures temperature and salinity. considered to be estuarine or coastal, while the naked forms are found more in the plankton of the open seas. Gymnodinium breve is apparently not estu- arine, but neritic. In controlled experiments, Aldrich and Wilson (1960) found the best growth at between 27 and 37 p.p.t. (parts per thousand). Field observations indi- cate ranges of salinity for good growth be- tween 31 and 37 p.p.t. From computer analysis of the 1954-61 field data we find no real abundance of the red-tide organism below 30.5 p.p.t. salinity (fig. 2). The upper limit for abundance of organisms appears to vary with temperature. The upper limit is 37 p.p.t.; growth is poor above 36 p.p.t, when tempera- tures exceed about 23° C.

Rainfall

Figure 3. in In Heavy rainfall is often given as a likely — Red-tide years relation to rainfall south factor in producing red tides. Thus, Hornell Florida, March-September, 1910-60. (1917) mentioned the annual occurrence of red tide "after the passing of the rainy season" that an outbreak will occur are enhanced by along the Malabar Coast. A translation of high rainfall. Thus, high rainfall must be Niimann (1957) follows: "According to all considered as one of the factors favoring reports, outbreaks of red water most obviously outbreaks. occur in coastal areas a short time after precipitation. The bloom of Exuviella occurred Wind in the coastal area of Angola after heavy rain- fall in the hills of Biniienland. The Congo and The effect of wind, or lack of wind, is often Cuanza Rivers brought nnuch water which was mentioned in discussion of factors favorable spread throughout the surface layers of for blooms. Thus, the spectacular blooms of coastal waters." On the other hand, red tide the diatom Aulacodiscus kittoni at Copalis occurs in many regions, expecially in areas Beach (Becking, Tolman, McMillin, Field, of upwelling, without abnormal rainfall. and Hashimoto, 1927) occur when a heavy- For the red tide in Florida the evidence rainstorm is followed by gentle westerly is somewhat contradictory. Thus, the heavy (onshore) winds. Blooms of Noctiluca nniliaris red tide of 1946-47 commenced in November on the Malabar Coast occur between monsoons 1946 after a comparatively dry spell and during calm weather (Bhimachar and George, ceased in September 1947 during very heavy 1950; Hornell, 1917; Hornell and Nayudu, rains. Because all eight recent outbreaks 1923). Blooms of Mesodinium rubrum in British commenced in the fall (table 2), we have Columbia occurred also during calm weather plotted the accumulated inches of rainfall for (Clemens, 1935). Lackey and Hynes (1955) 22 south Florida weather stations for March commented on the effect of heavy weather to September, inclusive. Tabulated data were in breaking up dense concentrations of G^. breve , available for 51 years, from 1910 to 1960 Pomeroy, Haskin, and Ragotskie (1956) com- (see app. table). mented on blooms of Amphidinium fusiforme The chance for an outbreak of red tide and Gymnodinium splendens , "All the Delaware appears to be much better in the autumn of Bay blooms occurred during periods of light years with heavy spring and sumnner rainfall winds, not exceeding Beaufort force 2 [4 to 6 than in other years (fig. 3). Only two red-tide knots]. On one occasion a patch disappeared outbreaks occurred in years with less than when the wind rose from force 2 to 3 [7 to 10 40 inches. The 1916 outbreak, although appar- knots]." ently heavy, lasted only 2 months (October and and was confined to the area November) Light from Boca Grande Pass to Big Marco Pass. Likewise, the 1952 outbreak commenced in Since most of the overblooming plankton November, stopped in January, and was con- that causes red tide is autotrophic, or depends fined to the areas adjacent to Sanibel Island, on organisms that are, light intensity must The other outbreaks were all more wide- be considered. Brongersma-Sanders (1957) spread, occurring at various points between stated, for instance, that plenty of sunshine Tampa Bay and Cape Romano. The innplication seems to be another requirement for red tide, is that rainfall alone has a rather low predic- Clemens (1935) noted that the bloom of Mesodi- tive value, but, at the same time, the chances nium in was preceded by "A couple of weeks of bright, sunny, calm and phosphorus. He stated that this concept weather." Conover (1954) listed "high radia- stems largely from observations that dino- tion values" as one of the essentials for a flagellate maxima, in temperate waters, fol- bloom of Gonyaulax . Hornell (1917) stated low the decline of the spring diatom flowering, that red tide on the Malabar Coast required and that relatively large populations often "A continuance of fine weather for a week or persist throughout the summer, when the ten days, with plenty of sunshine." supplies of these nutrients are alnnost unde- There is evidence, however, that light is tectable. seldom a limiting factor in G. breve abundance. Niimann (1957) appeared to hold similar Wilson (1955) obtained good growth of cultures views. He stated, "It also appears--and this by using 175 ft.-c. (foot-candles) for 15 hours is our final conclusion--that a mass outburst per day, and Aldrich (1960) stated that light of occurs when fresh-water intensity is probably not a growth-limiting growth-promoting substances (trace elements, factor above the 200 ft.-c. level. Aldrich (1962) enzymes, or other biologically active sub- established, however, that G. breve is auto- stances) reach the sea. Due to the presence trophic, requiring light for survival. of a necessary quantity of nutrients in the sea, a pre-condition for the outburst of plankton in the sea exists already. Accordingly, Upwelling these outbursts occur only near the coast." In many of the regions where red tides In studying the phosphorus values within and occur, nutrients are brought to the surface without red-tide water from the work of by upwelling of nutrient- rich waters. The several authors, Bein (1957) concluded con- nutrients do not of themselves appear to be cerning G. breve that, "It is possible that the the cause of overblooming, however. All threshold level of total phosphorus necessary authors seem to agree that red tides usually to support dense populations of this organism occur after the cessation of upwelling. For is lower than originally assumed. ... It seems instance, Brongersma-Sanders (1948) stated very probable that, insofar as phosphorus is that blooms of Noctiluca in Walvis Bay occur concerned, the areas of the west coast of during periods of minimum upwelling; she Florida which have recorded Red Tides are, also said (1957) that "in areas where upwelling at all times, capable of supporting an out- ." occurs during part of the year only, red water break. . . usually develops toward the end or directly It also appears that G. breve does not re- after the period of upwelling." quire much nitrogen. Lackey and Hynes (1955) In regard to the situation on the west coast stated that Howard Odum failed to find any of Florida, Graham, Amison, and Marvin nitrogen in a series of about 15 field (1954) showed that the surface waters of the samples from red-tide water, taken October 6, Gulf gradually decrease in phosphorus out to 1953. Dragovich (1960b) stated, "No relation- a distance of about 85 miles, and that limited ship was observed between the incidence of upwelling of deep water at certain times had G. breve and nitrate-nitrites." no apparent effect on the phosphorus content In considering the nutrients available, one of water in the euphotic zone. must note that the rivers of the Florida west coast- -especially those draining phosphatic formations --contain heavy concentrations of Nutrient Levels phosphorus (Odum, 1953; Specht, 1950; Graham It appears to be the concensus of many et al., 1954; Dragovich and May, 1962). After authors that red tides caused by dinoflagellates weighting the discharges of seven rivers do not depend upon a high concentration of according to relative concentration of phos- nutrients. In fact, dinoflagellates usxially bloom phorus, which differ considerably among the after the have imipoverished the water. rivers, we estimated the metric tons of phos- Thus, Brongersma-Sanders (1957) said, "The phorus discharged by them. The results are greatest outbreaks of red water probably shown in figure 4. The Tampa Basin rivers occur toward the end of a phytoplankton are the Hillsborough, Alafia, , and season. . . ." Hornel and Nayudu (1923), ac- Little Manatee; the south Florida rivers are cording to Ryther (1955), described red water the Peace, Myakka, and Caloosahatchee. The caused by peridineans occurring annually along picture does not vary too markedly from that the Malabar Coast, following the cessation of of rainfall. From these data one cannot assume the southwest monsoon, diatom blooms, and that phosphorus necessarily had any more heavy rainfall. Blooms of Gymnodinium sp. effect than another ingredient of river water. occur annually along the TrivandrumCoast of India, following rains and diatoin maxima, Are Preceding Diatom Blooms Conducive to according to Menon (1945). Red Tide? In summing up this reasoning, Ryther (1955) wrote that dinoflagellates have often been It seems to be fairly well established that credited with the ability to utilize and flourish red-tide organisms can and do bloom when in extremely low concentrations of nitrogen nutrient levels are low. There has been 1 Gunter (1952) reported, "In the summer as a detoxicating agent in cultures of

of 1935 a very heavy mortality of marine Prymnesiunn parvum . fishes and other animals occurred in the Since Aldrich (1962) has established that and extended for a distance G. breve is autotrophic, it would seem that of 250 miles from the Rio Grande northward. there would necessarily be some delay before A few fishes were killed in the lower bays the constituents of whole fish had had time but practically all of the mortality was con- to recycle inorganic nutrients into the water. fined to the Gulf. ..." In a later outbreak It should be noted that during the period in this area, Wilson and Ray (1956) identified of time necessary for putrefaction, surface-

the causative organism as Gymnodinium breve . drifting fish may be blown far offshore, or Semicircular surface structures were re- onto some beach, many miles from the red ported by Ingle, Hutton, Shafer, ajnd Goss water which killed them. Hela (1955) showed (1959) to be comnnon features of the 1957 that the drift of the dead fish can for practical red-tide outbreak near the mouths of passes. purposes be determined by wind observations They said: "In every case observed by the alone. Speaking of the 1947 outbreak, Anderson

authors (R.M.I, and R.F.H.) the discolored (USFWS, 1958) said,". . . in that summer some water, denoting a high concentration of dino- of the fish were pushed out, but that doesn't flagellates, was found from the interface and mean that you don't have offshore kills. The extending seaward." Ingle also stated (USFWS, big concentrations were inshore. Fish that 1958), "We made one trip along interphase and were far offshore were in clear water. Most found high concentrations as far as 15 miles fish found offshore were in a bad state of offshore. Very high counts as far as we could decay, which nnade me think they had been go." drifting for a long time after being killed Slobodkin (1953) based his hypothesis of in red water." Wilson (USFWS, 1958), when stable water masses wholly on theory. His questioned, answered that in general fish theory that organisms develop in a "discrete kills occur close to shore, except in certain mass of water of relatively low salinity" instances, and that he did not know of a bloom differs from observations on blooms occurring completely offshore, not associated with aun in streaks, the abrupt margins of the blooms, inshore bloom. In November 1954 during a and the formation of blooms at interfaces. period of strong northeast winds, the Oregon The theory does not explain howthe organisms, found dead fish 50 miles offshore. Finucane

floating in their lens of water, concentrate (USFWS, 1958) said, ". . . In 1947 I don't high announts of nutrients from nutrient-poor think they died near the Keys but were moved water. down by wind action."

Effect of Dead Fishes in Continuing a Red-Tide Bloom FIELD OBSERVATIONS ON FLORIDA The question of whether or not fish killed RED TIDE by red tide aid in perpetuating or spreading red tide is of some importance. Proponents To aid in guiding our thinking, we compiled of such an hypothesis point out the immense table 1, showing the occurrence of recorded quantity of nutrients that can be made avail- outbreaks of red tide during the 117 years able by the decaying carcasses. from 1844 to I960. We found records of 20 Collier (1955) squeezed juices from freshly toxic red tides, 3 in Texas and 17 on the killed fish into a culture he described as west coast of Florida, all of which can reason- about 3 million organisnns per quart; over- ably be attributed to blooms of Gymnodinium

night they tripled in number. Wilson (1955) breve . In evaluating these records, it should discovered, however, that if fish autolysate be borne in mind that, especially in the earlier is added to the original culture medium, reports, the observers did not separate cause bacterial growth frequently is stimulated, and effect. Reports of great quantities of and the culture dies. For instance, when fish drifting ashore at Key West or on the Dry added to a culture of 3 1/2 million per liter, Tortugas for instance, gave no indication of the organisms increased in 4 days to 9 1/2 the area or areas in which they were killed. million but then fell to zero in 1 week. Some of the recorded outbreaks may have In considering these results, it should be been minor. For instance, concerning the 1952 noted that when fish are killed by red tide, outbreak. Lackey and Hynes (1955) wrote, the fresh juices are not squeezed into the "The 1952 outbreak was apparently rather water; the fish undergo a process of putrefac- limited in the area affected, which seemed tion during which the bacteria nnust increase to center around Sanibel Island. There is enormously. In this regard Ray and Wilson little published information on this outbreak, (1957) mentioned the effect of bacteria in at least at the present time." Both Collier reducing the toxicity of G^. breve cultures, (1954) and Chew (1953) confirmed the oc- and also mentioned the experiments of Shilo currence of this outbreak, however, placing and Aschner (1953) in which bacteria acted it around the Sanibel Island area in November. Table 1. —Occurrences of Florida red tide, 18^ to 1960

Months No. General area Initial month Final month unknown

1- 1844 ' n. (^)... (^) 2. (1)... 1854 (J) 3. 1856 or '57 (^)... (') A. 1865 {'> 5. 1878 Tampa to Dry Tortugas Sept. (2). Oct. (1,5) Nov...... 6. 1879 7. 1880 Tampa to Key West Aug. n. Dec. (^). Egmont Key Oct. {'). 8. 1882 Clearvirater to Egmont Key July ('). 9. 1883 (8,9) 1884 10. 1885 E^mont Key to Charlotte Harbor. Oct. (^°). •fli-)-- 11. 1908 («) Grande to Big (faroo Pass.. Oct. (8).... Nov. (^)... 12. 1916 Boca "("*)" 13. 1935 South Texas June (^2,13). Aug. (12,") 14. 1946 Boca Grande and South Nov. (^')... 1947 Jan. (',1') 1947 Reached Tampa Bay in July Mar. Sept. ("). 15. 194e South Texas Fall (")... 16. 1952 Near Sanibel Nov. C-S,!'). 1953 Jan.(i''i8,i') 17. 1953 Tampa and South Sept. (18,19) 1954 Mar!""(20)!i! 1954 Venice and South July (21). 1955 Feb. (21)... 18. 1955 South Texas Sept. ("). Sept. (")... 19. 1957 Tampa and south Sept. (21). Deo. (^^)... 20. 1959 Tampa to Venice Sept. ("). Nov. (")... 1960 £gmont Key Mar. ("). Mar. (")... 1960 E^ont Key Juljr ("). Aug.

1 Ingersoll, 1882 ' Gunter, 1947 1' Univ. Miami, Mar. Lab., 1954 2 Jefferson et al., 1879 1° Glennan, 1887 18 Collier, 1954 ^ Glazier, 1882 11 Canova, 1885 1' Chew, 1953 12 ' Jefferson, 1879 Umd, 1935 2° Lackey and Hynes, 1955 21 ' Moore, 1882 1^ Burr, 1945 Finucane, 1964 * Walker, 1884 1" Gunter, 1952 22 Wilson and Ray, 1956 ' 1* 2^ Anon. , 1883 Gunter et al., 1948 Finucane, 1960 ' Taylor, 1917 1^ Galtsoff, 1948 2* Finucane, 1961

Seasonal Occurrence of Red Tide on temperature, nnentioned previously, which indicate that G. breve cam thrive at tenn. Table 2 (from data of table 1) shows the peratures from 16° to 27° C. This seasonal seasonal occurrence of red tides in Florida blooming in early autumn, however, could for the eight outbreaks for which the data also be encouraged by seasonal rainfall, which appear to be reasonably accurate as to month. is normally much higher in June, July, August, Several features are of interest. All outbreaks and September than in other months. commenced between August and November, and half started in Septennber. Six of the out- The Role of Convergences in Causing breaks ended between November and Febru- Red-Tide Blooms ary, suggesting the effect of lowered tem- perature in controlling growthof the organism. Several cogent arguments support the hy- At least three of the outbreaks apparently pothesis that convergences play an important did not cease during winter aind spring, but role in initiating cind maintaining toxic con- merely continued at a slow pace until more centrations of the Florida red tide. Some of temperate weather returned. these arguments are: Another interesting feature of table 2 is 1. The Florida red tide apparently can that of 48 months in which toxic blooms aggregate in fish-killing concentrations at were present, only 11 occurred during the a very low nutrient level; however, the phos- 6-nnonth period from February through July. phorus values within such concentrations (in- This coincides with the observations and cluding the organisms in the samples) are opinions of several authors that most dino- often high- -often far too high to be accounted flagellates bloom after the diatom flowering for without supposing the organisms to be and after the nutrients have become rela- capable of concentrating practically all of tively scarce. It also coincides with the data the phosphorus available in the . Table 2. — Seasonal occurrence of Florida red tides

Year 8. Despite strong tidal movements through never very abundant from March through the passes, and a general residual current August. For the 3-month period from Sep- flowing northward along the coast, a fish- tember through November, the contour lines killing concentration may remain in virtually of relative abundance are shown in figures the same place over several days at a time. 5 and 6. Centers of high abundance were in This persistence could be explained by the the Sanibel Island area and adjacent to the organisms being held in one area by a con- mouth of Tampa Bay. Heavy concentrations vergence at the mouth of the pass. Wilson tended to hug the coastline. A light concen- (USFWS, 1958), in discussing an early con- tration extending seaward off Tampa Bay may trol experiment, said, ". . .this bloom had be a reflection of the permanent counter- persisted in this area [0.2 square mile] for clockwise eddy which, according to Hela, de 22 days." He also stated, "If you find a high Sylva, and Carpenter (1955), seems to have concentration of organisms one day, it tends its northern end at the latitude of Tampa Bay. to be at this location the next day. If there is During 2 months (December-January) the water movennent you would expect to find them distribution was in general similar (figs. 7 at some other location and not at approxi- and 8), but the level of abundance had fallen mately the same position day after day." in the north, which may well reflect the There is some evidence that dinoflagellates curbing effect of cooler water. tend to sink toward the bottom at night and rise to the surface during the day. The Pollution as a Contribution to Red-Tide sinking may not be purposeful movement, Outbreaks but merely a tendency for the organisms to become dispersed at night when there is The question has been raised as to whether no light to bring them toward the surface. nutrient-rich pollution, especially domestic Pomeroy et al. (1956) mentioned that they , contributes in initiating or main- observed a bloom of Gymnodinium sp. that for taining outbreaks of red tide. Odum (1953) 4 successive days disappeared at night and found in Florida that 18 small unpolluted in the early nnorning and reappeared each streams contained 0.019 p. p.m. (parts per day at about 9 a.m. While spraying copper million) of phosphorus, 10 small acid streams sulphate during the control experiment in draining phosphatic formations contained 0.413 the heavy outbreak of 1957, it was noted p. p.m., and 7 streams not draining phosphatic that concentrations of G. breve were not formations but receiving sewage contained apparent in the early morning before the sun 0.836 p. p.m. The effects of sewage could attained a high angle. result from an increase in phosphorus and Because dinoflagellates cam move vertically nitrogen or from some other condition. Bac- they may be able in calm weather when con- teria working on sewage, for instance, might vergences are fewer to strain the nutrients increase the amount of VitaminBi2 available. from a goodly part of the water column. Wilson (USFWS, 1958) said ". . . possibly the That a dense concentration of organisnns is most important aspect of stream discharges usually formed either at a convergence or in which are contributing to red tides is that very calm water is shown by the fact that some form of chelating agent may be in this ." concentrations of G. breve often occur in water. . . places protected from currents, such as small If red tides are tending to occur more bays, or the stagnant fingers made by dredging frequently, and especially if they are tending narrow blind channels to make waterfront to occur more toward the northern portion lots (Ingle et al., 1959). Wilson (USFWS, of the red tide area, one might well suspect 1958) said, "You normally have more plankton that pollution from the mushrooming cities bloom and red tide during periods of calm around Tampa Bay is contributing to their weather." occurrence.

Coastwise and Seasonal Abundance LIFE FORMS OF GYMNODINIUM BREVE One of the first steps in analyzing the 1954-60 raw data on G. breve was to deter- Very little has been published on life forms mine how the organisms were distributed of G^. breve . Detailed descriptions of the both seasonally and in relation to the coast. organism appear in Davis (1948) and Lackey For this purpose the data were coded into and Hynes (1955). The latter stated that the 10 north-to-south areas, each measuring 25 only observed method of reproduction is by miles along the general trend of the coast binary fission. Wilson (USFWS, 1958) men- (fig. 1). To obtain the distribution in relation tioned observations on cysts. He stated that to the land, the data were also coded in he could not make G. breve encyst by tem- bands of distance offshore, and, for stations perature adjustments. When questioned con- inside the fringing barrier islands, by dis- cerning whether cysts would divide, he stated, tance to the mouth of the nearest pass. It "I have put in from one to 20 in sterile media was obvious that in 1954-60 organisms were and found that they will encyst and divide." He

10 also stated, ". . . Cysts form chains and when three main aspects, 1) effect of red tide on culture is stirred these tend to remain to- the tourist business, 2) effect on the sport gether, but I have never seen a chain for- and commercial fisheries, and 3) effect on mation of normal free swimming cells." public health. There are several statennents that G, breve affects the viscosity of the water when nu- Damage from Red-tide Outbreaks merous. Galtsoff (1948) stated, ". . . the water had an oily appearance. When dipped up and Since red-tide outbreaks usually start in allowed to stand for 5 to 10 minutes, it be- the fall, just as the tourists are commencing came thick, sometimes almost of a consistency to move south, an outbresik can cause very ." of Karo syrup, and slimy to the touch. . . serious losses, especially at seaside com- Gunter, Smith, and Williams (1947) reported, munities in which the accommodation of "The water was viscid and slimy, having the tourists is often the chief, or son-ietimes consistency of diluted syrup." Collier (USFWS, the only, major business. Rotting fish on

1958) said, ". . . observing with a dissecting the beaches, cind sometimes acrid aerosols microscope, the viscosity seenned to be caused containing , will drive tourists away. by the organisms agglutinating. While I watched From this standpoint alone, control is highly they started joining in long chains, long desirable. branching chains. ..." The effect on the fish populations has not There may be another answer to the high been nearly as severe as the layman imagines viscosity. Martin and Nelson (1929) found when he hears of the death of millions of small by killing Gynnnodinium sp. with special solu- fishes. The percentage kill is doubtless low. tions that they could observe a gelatinous For instance. Lackey and Hynes (1955) stated, envelope as thick as the dianneter of the cell "... the last localized outbreaiks of the 1953- itself. They stated, "Gelatinous envelopes 54 Red Tide are only a few weeks past, yet are common among dinoflagellates when en- sports fishing (grouper, speckled trout, cysted, but not when active. The cells re- mackerel, and redfish) has been generally ." ferred to in this connection were actively good throughout the entire area. . . nnotile. A similar envelope has been noted The kill of fishes by the 1946-47 red tide occasionally surrounding other species of was estimated by Gunter, Williams, Davis, naked dinoflagellates in the active condition, and Smith (1948) as 500 million fish. This but only when killed by the iodine or bi- figure nnay sound large, but actually it is not. chloride methods. In red water plankton in If we realize that these fish will probably which Amphidinium fusiforme is the dominant run no less than 10 to the pound, the total species, the Amphidinium cells tend to cling is only 50 million pounds. Every year about together in clunnps, but no gelatinous envelopes 1 billion pounds of are caught along can be demonstrated. In many of the clumps a 300-mile stretch of the northern Gulf, but (although not in all), however, they may be this enormous catch of a single species does seen to be clustered thickly about a cell of not appear to be harming the supply. the Gymnodinium . This gelatinous envelope The effect of red tide on public health is may well be a factor of importance in hold- rather debatable. For a short while during ing the organisms together, once they are severe outbreaks there will be aerosols from massed by a favorable combination of light, breaking surf when the wind is blowing on- water temperature, and tidal currents." shore. Long exposure close to, or on, the Conjugation of G. breve was discussed by beach can be very irritating to the respiratory Wilson (USFWS, 1958). He said, "In the con- tract. A few people are so adversely affected dition I have called conjugation I have never that they must leave until the outbreak is seen more than 2 orgcinisms attached together. over. More serious have been the questions You may find them in various positions, but regarding the safety of products, there is always a stalk-like process between shellfish in particular, during an outbreak; two individuals." however, the Gulf Coast Shellfish Sainitation The size of G. breve is usually given as Research Center (1964) stated that, "... rig- between 20 and 30 1^ . Wilson (USFWS, 1958) orous proof of the Red Tide organisnn as

said, ". . . In certain cases in the field you the cause of shellfish toxicity remains to be will see a fairly large form which appears demonstrated." Since Gonyaulax and other ." to have no chromatin material at all. . . dinoflagellates occur, often in fair abundaince, in the red-tide area of the coast, it seems hardly fair even to attempt to indict G. breve CONTROL OF RED TIDE without very definite proof. tend to occur, except when trauisplauited, in areas It is difficult to discuss control of red tide with salinities belowthose tolerated by G. breve without first discussing the type and scope (which is neritic), but well tolerated by several of the damage inflicted. This damage has species of estuarine dinoflagellates.

12 Methods of Control suitable for blooms, or to suddenly lower the inshore salinity below tolerable levels Methods of control of red tide depend for G. breve should be considered. One might largely on whether the objective is preven- speculate as to whether the former larger tion, control, or nnere alleviation. flows of southward through the Under alleviation fall such measures as had any effect on earlier red seining of dead fish to keep them off the tides, which seem to have been more southerly beaches; also, the common practice of using in occurrence. bulldozers and other power equipment to bury 4. Encouragement of competing or inhibi- mechanically, or to gather together amd re- tory organisms by special types of fertiliza-

nnove, the dead fish washed or blown ashore. tion . This idea appears to be innplicit in the At best these are temporizing measures. As suggestion of the University of Miami Marine we have noted previously, fish-killing con- Laboratory (1954) that the carrying out of centrations of red tide tend to remain close extensive fish culture in the back bays might to shore, sometimes inside the passes. These cause changes in the habitat detrimental to

temporizing measures do not help against G. breve . The danger here lies in losing the the irritating aerosols, nor can they keep estuarine nursery areas needed by shrimp all fish off the beaches when the concentra- and various sport and commercial fishes. tions innpinge so closely on the fringing 5. Altering the physical habitat through con- islands. Numerous suggestions have been made struction of underwater barriers, jetties, or for both prevention and control, each of which similar structures at passes to restrict mixing we discuss in turn. of Gulf cind bay waters, and perhaps to change the pattern of the convergences at the mouths

of the passes . This suggestion (which we Prevention of Outbreaks have slightly elaborated) was made by Robert Prevention of outbreaks before they get Hutton at the meeting of the Advisory Com- started involves alteration of the habitat either mittee at the 1958 symposium (USFWS, 1958). chemically or physically. It certainly deserves full attention. 1. Inhibiting the growth of G. breve by raising and nnaintaining the concentration of Control of Outbreaks heavy metals at an inhibitory level . The idea here was to raise the level of copper (or Only a few of either the preventive meas- other heavy metal) in the inshore waters ures listed above or the control measures sufficiently high to inhibit the growth of the discussed below have been tried, even on a red-tide organism without actually destroying laboratory scale. Nevertheless, at the present it. Should an outbreak nevertheless occur, state of our knowledge, we cannot afford to only minimal amounts of copper added to dismiss or reject ideas in cavalier fashion. the water should suffice to kill G. breve Sonne of the untried suggestions undoubtedly because of the existing high level of copper. have merit- -it is a question of patient re- The feasibility of this idea was tested by search, pilot experiments, and economics. using copper ore, and the method was re- 1. Biological control through the use of jected (see Marvin 1958, 1959; Marvin, Lans- bacteria . The use of bacteria to destroy the ford, and Wheeler, 196l). toxin of the red tide was suggested at the 2. Keeping the nutrient level low by curbing 1958 symposium (USFWS, 1958). The use of use of streams zmd bays for disposal of bacteria that destroy vitamin Bj2 was sug- nutrient-rich sewage or other pollutants . Since gested by Hutner and McLaughlin (1958), but G. breve Ccui apparently thrive in nutrient- dismissed with the comment that "truly poor water, this suggestion does not appear enormous quantities of bacteria would be ." too promising. We do need to know whether required. . . sewage pollution contributes other substances 2. Biological control through encourage- needed by the organism. ment of predator organisms . The encourage- 3. Controlling river flows in streams for ment of predator organisms was mentioned which water can be stored so as not to create by several authors. Galtsoff (1948) mentioned

ideal conditions for blooming . Some streanns, the ingestion of G^. breve by a cladoceraui, particularly the Caloosahatchee River, have Evadne. Hutner and McLaughlin (1958) men- enormous storage capacity. By regulation of tioned the protozoans amd the luminous the flow (a detailed hydrographic study at all dinoflagellate, Noctiluca. Torrey (1902) men- flow levels is highly desirable) it may be tioned the appearance of Noctiluca in great possible to prevent the formation of water nun-ibers toward the end of July amd their of suitable salinity in a zone of strong con- devouring Gonyaulax "with avidity." There vergences. Seasonal releases of blocks of is no comnnent on how to encourage these water, either to get rid of surplus water predators, short of providing them a red- during a period when conditions appear un- tide bloonn.

13 3. Physical control by high-frequency In a large-scale experiment, during the waves . Little attention has been paid to physi- heavy outbreak of 1957, 105 tons of copper cal means of control. Lackey and Hynes (1955) sulfate were spread by crop-dusting planes stated, ". . . It has already been shown, how- at about 20 pounds to the acre along 32 miles ever, that there is no useful killing action in of beach between Pass-a-Grille and Anclote ." a high-frequency radio field. . . Keys (Rounsefell and Evans, 1958). 4. Control by changing the pH. Galtsoff 7. Destruction of G. breve by selective

(1948) wrote ". . . The use of powdered calcium chemicals . The primary objective is to dis- oxide (unslackened lime) suggests itself, for its cover a chemical or chemicals toxic to G. breve addition to sea water will raise the pH to a at low enough concentrations to hold promise level which is beyond the tolerance of the of being cheap enough to use for control, ." dinoflagellate. . . [p. 35.] Wilson (1955) said that at the same time is specific for G. breve that G. breve grows best in pH of 7.3 to 8.1, or perhaps closely related species. The but survives pH from 7.0 to 8.6. Aldrich (1 959) screening of 4,306 compounds, including most gave 7.5 to 8.3 as the pH range for good of those screened in past years in developing growth of G. breve and said that a pH below a specific larvicide for the sea lamprey

7.3 was definitely toxic. ( Petromyzon marinus ) in the Great Lakes, 5. Control by adsorption of vitamins or yielded 55 compounds toxic at 0,01 p. p.m. chelators or both . Adsorption of vitamins (Marvin and Proctor, 1964). Further tests required by G. breve through large-scale (unpublished) have shown that several of these dusting with charcoal was suggested by Odunn, highly toxic compounds do not harm several Lackey, Hynes, and Marshall (1955) as a other estuarine organisms. The laboratory nneans of modifying offshore blooms. This testing of these very toxic compounds, when suggestion should be tested in the laboratory. completed, should lead to pilot experiments 6. Destruction of G. breve by nonselective under field conditions. chemicals . The most widely tried method of control for red tides has been the spreading or dusting of water areas with nonselective SUGGESTIONS FOR FUTURE RESEARCH chemicals. The question of what destruction In summarizing research to date on red these chemicals can do to organisms other tide, we are immediately aware of the fact than the one in bloom is usually based on that, despite some very significant progress, general uneasiness, without substantial proof. many gaps remain in our knowledge. When For instance, few question the spraying of we consider the tremendous expenditures of valuable estuarine nursery grounds withprac- time and effort that have gone into the solu- tically nondestructible hydrocarbons in the tion of many medical problems, such gaps name of mosquito control, yet many worry are not surprising. Serious red-tide research about a very low concentration of copper is comparatively recent; barely 10 years have sulfate, which has only a momentary effect. elapsed since the first successful culturing Several nonselective chemicals have been of the causative organism. Perhaps the chief suggested for killing red-tide organisms di- deterrent to progress has been the fluctuation rectly: in financial support. Heaviest support has Ammonia was sharply inhibitory to growth followed severe outbreaks; support has de- of ; it was less toxic at clined between outbreaks until, at times, the high salinity or at low pH (McLaughlin, 1958). level has become too low to provide ade- Ferric chloride and chlorine gas were used quate continuity of field data. Despite the in Japan; materials were discharged over marked improvement in the field programs the stern of a boat so they could be churned since the 1958 symposium finances have been into the water by the propeller (Anon., 1934). insufficient to keep up continuity in field and Calcium hypochlorite or liquid chlorine laboratory and at the same time carry out was used by the Japanese either after or research on some of the imaginative sug- with copper sulfate to prevent the growth of gestions that were made. Perhaps some re- bacteria after killing the dinoflagellates orientation may aid in stretching the research (Galtsoff, 1948). dollar. Copper sulfate is the most widely used Several items suggested by the Advisory chemical. The Japanese have used it for Committee at the 1958 symposium have been many years to protect their beds worked on, and some completed. These items from red tide. It was tried on a small scale follow: in Florida in 1952 by discharging a concen- 1. Testing chemical comp oTindg to discover trated solution of 3,000 pounds of copper sul- one or more specific for G. breve . This work fate from the ballast tanks of the Alaska . In has resulted in a report by Marvin 2uid Proc- 1953 sacks of copper sulphate were towed fronn tor (1964) for 4,306 chemicals, several of small boats off Anna Maria Key. Later a small which show promise. This item should be area was dusted by plane. None of these ex- pushed vigorously. periments was on a sufficiently large scale to 2. Does G. breve require a heterotrophic indicate the effectiveness of copper sulfate. existence? Aldrich (1962) showed that G. breve

14 is photoautotrophic, requiring both light and favoring resumption of the normal form. The CO2 for growth and survival. winter survival of G. breve in the deeper 3. Would a copper ore dike maintain a (and warmer) offshore waters (as deep as high enough level of copper in the water to 123 feet) has been stressed (Dragovich, 1960b; inhibit growth of G. breve ? The answer is Dragovich and May, 1961; Finucane, 1960), no (Marvin, Lansford, and Wheeler, 196l). but overwinter survival of encysted forms The Advisory Connmittee urged the Bureau in the shallower inshore waters is a dis- of Commercial Fisheries to attempt to pub- tinct possibility that requires investigation. lish both the data and the results of its red- Should this inshore survival occur it nnight tide investigations. Since 1958 the Bureau has be remotely possible, following heavy red- published all its raw field data and a number of tide outbreaks in the fall, to modify the sudden reports on both field and laboratory projects. recurrences in the spring (as happened in 1947, 1954, and I960) by destruction of the cysts on the bottom. Laboratory Studies of the Organism Binary fission was the only method of repro- Laboratory studies of both unialgal amd duction observed by Lackey and Hynes (1955). bacteria-free cultures have yielded consider- Conjugation was also observed by Wilson able information on the physiology of G . breve . (USFWS, 1958). Because it is apparently not More work is needed, however, especially often observed, the discovery of what favors on the role of other plankton organisms in conjugation and its role in maintaining popula- promoting or inhibiting growth. This type of tion abundance might be important. work may involve the use of radioisotopes Although the red-tide organism normally to trace the flow of nutrients. measures about 20 to 35 (i , large forms, per- There has been much discussion of the haps up to 80(1, have been observed. Wilson possible role of chelators in stream water (USFWS, 1958) mentioned that occasionally or surface runoff in promoting growth. in the field one observes "... a fairly large In both of the above cases it is possible form which appears to have no chromatin that experiments have been hampered by the material at all, . . ," He did not observe very high surface:volunne ratio of the test these in the laboratory. It would be of in- tubes in which most experiments have been te rest- -perhaps of some importance --to dis- carried out. In 1958 several advisers sug- cover whether this large fornn is a special gested that cultures be grown in large tanks life form of G. breve or perhaps a distinct to answer better such questions as killing species. concentrations of G. breve and the effect of Opinions seem to differ as to whether dead fish, either in perpetuating a bloom, G. breve csin form chains. Wilson (USFWS, or possibly in creating lethal concentrations 1958) said that cysts form chains, and that of bacteria. when the culture is stirred these tend to Some have advocated attempting to cause a remain together, but he had never seen a bloom of red tide by such measures as chain formation of normal free-swimming dumping a barge of dead fish, dumping large cells. Collier (USFWS, 1958), however, men- amounts of pollutants from selected localities, tioned watching the organisms form long or by fertilizing an area containing organisms chains. His observation was apparently made with specific nutrients. Most agreed that to while he was watching a very high natural gain sufficient control over the experiments, concentration--probably much higher than any the studies might better be performed in encountered in the laboratory. It is possible large tanks. Discovering how to create a that dense concentrations may cause the bloom is the opposite approach from ob- organisms to act in a different manner. The serving blooms in the field and then trying possibility that a difference in life forms to decide the cause. We believe the prob- or manner of reproduction may be triggered lem should be approached from both direc- by excessively high concentrations deserves tions. In any such tank experiments, we investigation. believe that an attempt should be made to There appears to be some question whether imitate natural conditions insofar as prac- changes in color of red-tide water are caused ticable. Thus, a very large tank might offer by density of the orgcinisms, by the angle of a fine opportunity to test the role of con- the light, or by the age of the population, vergences in concentrating the organisms, Wilson (USFWS, 1958) said that each culture and perhaps to solve their ability to grow seems to have a tendency to accumulate oil in water of low nutrient content. droplets with age, which will give a different The life forms of G. breve have been ob- color comprehension. If either color or oil served, but work is needed to determine the accumulation could be used as a gross meas- factors causing encystment, the conditions ure of population age, it might be useful in under which cysts can survive, the length determining whether a population is increasing of tinne they can survive, cuid the conditions or declining.

15 Field Studies of Red Tide favorable to G. breve , or 3) by providing competition, perhaps through excretion of a

Since 1954 the Bureau of Commercial Fish- substance detrimental to G. breve . eries has gathered environmental data that These quantitative data on other plankters, included concurrent enumerations of G. breve . thus, need to be supplemented by laboratory One of the chief aims when this sampling work on the effect of these plankters, or of began was to attempt to predict red-tide the water in which they have grown, on the outbreaks by noting the gradual increase in growth of G. breve . It nnay be necessary to abundance of the orgEinisms. Unfortunately, grow cultures of a number of plankters. high concentrations of the organism appear These cultures should include forms that with little prior warning. In 1957 this pro- prey directly upon G. breve . If predatory gram was revised to give nnuch more in- forms could be encouraged by any feasible tensive coverage of one area- -Tampa Bay means, this type of control would be pref- and vicinity. More chemical determinations erable to the use of nonspecific algicides. were also added. At the meeting of the Advisory Committee One of the first tasks should be to complete at the 1958 symposium (USFWS, 1958) Rice the analysis of this accumulation of environ- suggested that field observations be made in mental data. It is inriportauit to know which an area outside the areas in which red-tide factors can safely be dropped, which must be outbreaks normally occur. Comparison of retained, and how much sampling is needed chemical, physical, and biological data from for a reliable estimate of population changes. the two areas might serve to isolate one or In the analysis of these data we feel it im- more causative factors. portant to obtain detailed data on wind direc- tion and velocity. If convergences play as Control Methods important a role as is strongly suspected in the rapid concentration of organisms, and Control methods have been discussed at possibly equally innportant, in permitting the some length in the section "Control of out- organisms to grow in low-nutrient water, breaks." Here we discuss only new or un- then winds may be one of the major factors tried methods. in overblooming. Any modifications of the The chief deterrent to the use of non- present sampling program should await the specific algicides is the possibility of results of the analysis of the accumulated inflicting serious damage to other forms. data to prevent discontinuity of the records. If applied carefully in concentrations just

A detailed hydrographic survey of the areas high enough to kill G. breve , this damage adjacent to the nnouths of the principal passes will be chiefly to plankton, including larval should be made under varying conditions of forms. Since dinoflagellates tend to concen- tide, wind, and residual outflow of brackish trate in convergences because of their ability water. This survey should include an attempt to resist vertical currents, it might be feasible to locate the position of convergences amd to control the red-tide organisms by treat- the speed of the vertical currents. Once this ment of the convergences only and, thus, detailed information is available for a major inflict minimal damage to other forms. This pass we suggest a model study to determine end might be accomplished by gradually dif- whether it is feasible to make smy modifica- fusing the algicide from floating material, tions of the pass (such as underwater barriers such as coarse sawdust. to restrict mixing, jetties, etc.) to aid in two in fresh water lakes have been suc- objectives: 1) to lessen the formation of cessfully controlled by the use of aniline dye convergences and 2) to prevent the formation (Eicher, 1947). In a 200-acre lake averaging of large bodies of nnixed Gulf and bay waters 10 feet deep, nigrosine dye was applied at of salinities favorable for G_. breve. 10 pounds per acre. The water became black, One importamt aspect of the field work and 18 months later retained about 50 percent that may need nnore emphasis is obtaining of its dark color. One very noticeable effect reliable quamtitative data on the relative was the lowering of the pH. Whether it is abundance, both seasonally and annually, of feasible to shield the light by use of dye- other plankters. G. breve cam grow in water stuffs, or perhaps sonne cheaper material, of relatively high salinity but appears to sufficiently to affect G. breve significantly bloom more often in the fall, usually at lower should be investigated. salinities.-suggesting that the stimulation of growth depends upon some terrigenous sub- Need for Coordinated Research stance brought into the Gulf by the rivers, or upon previous chauiges in the neritic waters It has become abundantly clear that research dependent upon other plankters, or both. The on red tide involves several disciplines-- influence of the previous plankton population both physical and biological oceanography, could operate in at least three ways: l)through microbiology, ainalytical chemistry, bio- rennoval of some substance inhibitory to chemistry, aind mathematics; several types

G. breve , 2) by excretion of a substance of physical equipment and infinite patience also

16 are needed. Above all, a steady level of fund- ALDRICH, DAVID V. ing is required. Emergency funds are of slight 1958b. Toxicity of copper to marine orga- value, regardless of amount, compared to a nisms. In Annual report of the Gulf moderate but reliable budget. One cannot do Investigations for the year excellent basic research if he is required to ending June 30, 1958, U.S. Fish Wildl. fire investigators periodically and then hire Serv., p. 86. whatever personnel are unemployed when funds become available. "A series of copper toxicity studies So much work still is needed, despite the is being conducted to compare the sensi- fine work already completed, that duplication tivity to copper poisoning of several of effort should be avoided- -although in science organisms representative of the Galves- duplication is seldom wasteful because, by ton Island Lagoon. the very nature of research, no two people . . . "To date, the studies suggest that ever approach a problem in exactly the same several Lagoon organisms maybe ar- manner. We would recommend, however, that ranged in this order of decreasing sus- Federal, State, and university personnel ar- ceptibility to copper: Littorina, young range for periodic discussions and continue sciaenids, young sparids, young mullet, their spirit of friendly teamwork in the in- grass shrimp (embryonated eggs as terest of effective research. well as large emd small adults)."

REFERENCES PERTAINING TO RED TIDE ALDRICH, DAVID V. AND RELATED SUBJECTS 1959. Physiological studies of red tide. In Galveston Biological Laboratory fish- ABBOTT, B. C, and DOROTHY BALLANTINE. ery research for the year ending June 30, 1957. The toxin from Gymnodinium venefi- 1959, p. 69-71. U.S. Fish Wildl. Serv., cum Ballantine. J. Mar. Biol. Ass. U.K. Circ. 62. 36(1): 169-189. The toxin produced by dinoflagellates "A study of the effect of pH on growth amd causing paralytic shellfishpoisoning of G. breve in bacteria-free cultures may be different from that of other shows that growth is unhannpered by dinoflagellates associated with mass pH's of 7.5 to 8.3, inclusive. Growth mortality of fish. The authors were took place at a reduced rate at a pH of unable to render shellfish poisonous 7.3, and lower values were definitely with it and stated that Fingerman et al. toxic. Mediunn having a pH of 7.2 was (1953) indicated that the paralytic shell- 100 percent lethal to this organism fish operates in a curarelike within six days, while 7.0 killed all manner, while they showed that Gym- cells within two days. [p. 69.] nodinium toxin depolarizes the excitable "Over 800 individual cultures of membrane. G. breve have been exposed to specially controlled temperature conditions ALDRICH, DAVID V. .... the organism did not survive temper- 1958a. Histological techniques for Gym- atures of 7° C. and below, or 32° and nodinium spp. In Annual report of the above. Population growth did not occur Gulf Fishery Investigations for the year at 30° (or above) and survival was ending June 30, 1958, U.S. Fish Wildl. very poor. Multiplication was visibly Serv., p. 94. slowed at 15° C, but not completely "The fragility of G. breve is clearly halted. It is apparent that the level at indicated by results obtained with usual which temperature becomes absolutely histological fixatives. Formalin, limiting to growth lies between 7° and Bouin's, Zenker's, and Schaudinn's 15° C, but the work which will deter- fluids yielded only a very occasional mine this level more exactly is not yet recognizable cell. Alcohol, acetone, and complete. Optimal growth can occur trichloracetic acid also left few intact in a range of temperatures including organisms, and iodine vapor, Lugol's 20° to 27° C. ... [p. 69.] solution, and a mixture of chromic and "These results strongly suggest that acetic acids were only slightly better. temperature can be an important factor Best results have been obtained with in limiting the geographic distribution

0.5 cc. of 1% osnnic acid per 10 cc. of of G_. breve . In this regard it may also G. breve culture. This method will best be noted that Florida red tides occurring preserve the living appearance of the in the fall have ceased with the advent organism, yielding about 70 percent of of the first cold weather of the subse- the live cells in recognizable condition." quent winter." [p. 70.]

17 ALDRICH, DAVID V. ALDRICH, DAVID V. 1960. Physiology of the Florida red tide 1962. Photoautotrophy in Gymnodinium organism. In Galveston Biological Lab- breve Davis. Science 137(3534):988- oratory fishery research for the year 990. ending June 30, 1960, p. 40-42. U.S. Fish Wildl. Serv., Giro. 92. Extensive experiments failed to show evidence of heterotrophy. G. breve re- The red-tide "organism seems unable quires light and CO 2 for growth and to tolerate fresh or brackish water and survival. thus probably subsists offshore between red tides." [p. 40.] "If Florida west coast rivers do not The optimum laboratory temperature provide direct energy sources for mul- range for growth of pure G. breve tiplication of G. breve , the organism's culture is said to lie between 15° and vitamin, trace -metal, and chelator re- 30° G. ". . . it is apparent that G. breve quirements assume added ecological can tolerate slow temperature varia- significance as factors potentially limit- ." tions better than rapid ones. . . ." ing population growth. . . [p. 990,] [p. 40.] Parent stocks were grown at 25° C., and parental influence on survival of cul- ALDRICH, DAVID V., and WILLIAM B. tures at varying temperatures is ques- WILSON. tioned. Results of tests to determine I960. The effect of salinity on growth of parental influence were not conclusive. Gynnnodinium breve Davis. Biol. Bull.

"The culture work of W. B. Wilson (Woods Hole) 1 1 9(l):57-64. demonstrates G. breve to be capable of ." autotrophic nutrition. . . [p. 40.] Bacteria-free cultures were grown Seventeen carbohydrates were tested in test tubes in an artificial medium. and none were "found to support G. breve Tubes inoculated with 100 to 200 G. breve light. populations in the absence of . . . cells. Growth estimated by visual ex- At this early stage of the work there amination with microscope. Classified is no evidence that this organism can growth into 11 categories. Three top derive energy directly from amy source categories were arbitrarily defined as other than light." [p. 41.] "peak populations." Rough calibration by Varying light intensities showed that actual count showed peak population light intensity is probably not a growth- estimates to contain not less tham 750

limiting factor above the 200 ft. -c. (foot- cells per ml. smd usually from 1 to candle) level. several thousand per ml. Tubes ex- The organisms were destroyed under amined at 4, 10, 18, 25, and 35 days. ultraviolet light and showed a preference Seldom any growth after 35 days. Five for blue or green light. experiments each with 9 salinity levels, The organisms also seem to prefer and 10 tubes at each level. In first polarized light, and "cell counts to date experiment salinity ranged from 6.3 suggest that polarized light may stimu- to 41.1 p.p.t., in last four experiments late more rapid growth than ordinary from 22.5 to 46.0 p.p.t. Their sug- light." [p. 42.] gested optimum salinity range for good growth of 27 to 37 p.p.t. agrees well ALDRICH, DAVID V. with the computer analysis mentioned 1961. Culture and nutrition of Gymnodinium in the sumnnary.

breve . In Galveston Biological Labora- tory fishery researchfor the year ending ALLEN, W. E. June 30, 1961, p. 58. U.S. Fish Wildl. Serv., Circ. 129. 1933. "Red water" in La JoUa Bay in 1933. Science 78 (2010): 12-13.

". . . We have recently compared the growth of G. breve in water from several Discoloration to "muddy red" or "dirty rivers in red-tide and non-red-tide red" caused by P rorocentrunn areas. A connpletely defined artificial micans. sea water, incapable of supporting growth by itself, was employed as a ALLEN, W. E. diluent for the river waters. The Alafia, 1935. "Yellow water" in La Jolla Bay in Hillsborough, Peace, and Caloosa- 1935. Science 82(2127):325-326. hatchee Rivers of Florida and the Atchafalaya and Sabine Rivers of Louisi- Yellow water (apparently harmless) ana and Texas were tested. Growth oc- was caused by a small unidentified curred only with the addition of Hills- flagellate, colorless and bearing four borough or Peace River water." flagella.

18 ALLEN, W. E. "Sampling over a prearranged series 1943. "Red water" in La Jolla Bay in 1942. of stations extending from the rivers to Trans. Amer. Microscop. Soc. 62(3): the 100-fathom contour for a study of the 262-264. local oceanography, nutrients, and plankton, has been completed. Data col- ALLEN, W. E. lected are being analyzed. 1946. "Red water" in La Jolla Bay in 1945. concentrations in this area are in agree- Trans. Amer. Microscop. Soc. 65(2): ment with those found elsewhere in the 149-153. Gulf inside the 100-fathom contour. "All plankton collected during the ANDERSON, WILLIAM W. past year has been examined and data 1951. Gulf Fishery Investigations. In An- have been set up on punch cards for nual report for fiscal year 1951, Branch analytical studies, [p. 32.] of Fishery Biology, Fish Wildl. Serv., p. 31-33.

"Plankton collections have been made "Culture studies on nutritional re- by conventional methods and by means quirennents of dinoflagellates and other of a high speed sampler and a newly de- nnarine organisms have been continued vised continuous sampler. The latter at the Service's Beaufort, North Caro- methods are still in the experimental lina, Laboratory." [p. 33.] stage but results are pronnising. ANDERSON, WILLIAM W. 1952. Gulf Fishery Investigations. In An- nual report for the fiscal year 1952, "In comparison to areas like the Cali- Branch of Fishery Biology, Fish Wildl. fornia coast or the North Atlantic, Serv., p. 16-18. plankton in the Gulf is sparse. Tows in those areas produce 8 to 10 times more "Effort expended in a new laboratory plankton than comparable tows in the being set up for artificial culture of Gulf. It has become evident, too, that a marine willbe directed greater abundance of plankton exists toward understanding nutritional and en- over the continental shelf than exists in vironmental requirements for survival waters beyond the shelf. A preponder- and multiplication of various organisms, ance of fish eggs and larvae also has particularly dinoflagellates. Results will been found within the 100-fathom con- be used in producing mass cultures to tour. determine the role of various organisms "Analyses for inorganic phosphate and in production and utilization of various nitrate have been completed on a total organic compounds which are being of 371 samples obtained on the first isolated front sea water. These results three cruises. It has become evident in turn, will be applied to studies on fronn these analyses that phosphate and young fish survival. nitrate concentrations are extremely " Red tide research. --A theory has low at all levels inside the 100-fathom been formulated that 'red water' de- contour. In that portion of the Gulf out- pends on occurrence of isolated non- side the 100-fathom contour, extremely mixing water masses. A study of small- low concentrations of phosphate and scale 'red waters' in some inland bays nitrate exist in surface waters, but rise and information from various other steadily to a maximum between about 450 sources support this hypothesis." to 600 fathoms, below which their con- [p. 18.] centrations decrease slightly, [p. 31.] ANONYMOUS. 1881. Mortality of fish in the Gulf of Mexico. Ann, Mag. Natur. Hist. 5(8):238-240. "Results to date lend weight to the theory that, in general, waters of the This article cited from Hutton (1956) Gulf of Mexico beyond the 100-fathom apparently contains same information curve are relatively sterile. It may be as in Moore (1882). stated tentatively that the economy of our fisheries is closely associated with that portion of the Gulf lying inside the ANONYMOUS. 100-fathom contour, the inshore waters, 1883. Poisoned water in the Gulf of Mexico. and contiguous land areas. U.S. Fish Comm., Bull. 2:104. [Cited by Hutton (1956) as from the Sunland Tribune, Tampa, July 20, 1882.] 19 "We learn from Capt. William Jack- ANONYMOUS. son, of the steamer 'Lizzie Henderson,' 1956. The red tide--a progress report. that on his trip fronn Cedar Key, Tues- First Nat. Bank of Dunedin, Dunedin, day, he encountered a streak of poisoned Fla., Bull. 1(6):1-11. water, covered with all varieties of dead fish, of more than a mile in extent, "Similar occurrences of varying in- off Indian Pass, between Clear Water and tensity have been reported along Egmont Light. The captain says that a Florida's West Coast both before and very offensive smell arose from it, and after the outbreak of 1947. Earliest that a good many bottom fish, such as chronicle is by Capt. William Jackson. eels, were floating dead on the surface. On a trip from Cedar Key in 1842 he We opine that this fact upsets the theory encountered 'a streak of poisonous water of some as to this poisoned water being covered with all varieties of dead fish, fresh water from overflow on the main- of more than a mile extent, off Indian land, impregnated with poisoned vege- Pass, between Clearwater and Egmont table matter, as there are no streanns of light.' Fish kills were reported subse- any size flowing into the Gulf near quently in 1854, 1878, 1880, 1882, 1883, where these fish were found." 1885, 1908, 1916, 1932, 1946-47, 1951, 1952, 1953 and 1954. There have been ANONYMOUS. independent recurrences within less than 1934. Chemicals repulse ravages of "red one year of each other while periods of

tide." Sci. Amer. 15 1 (4):200-201. at least fourteen years have elapsed with no reported outbreak, [p. 3.]

"Turning back the 'red tide,' which periodically sweeps in from the sea to wreak havoc with the pearl-oyster beds "...There are plans this summer along the coast of Japan, is one of the [1956] for further study with visible nnost recent and romantic applications of tracing materials of the flow of river ." modern chemistry. Large areas of the discharge into the Gulf. . . [p. 11.] sea along the Ise Bay are used for the The 1842 date was doubtless 1882 cultivation of pearl-oysters and the (see Anonymous, 1883). jewels produced by the crustaceans are ARMSTRONG, F.A.J., and G. T. BOALCH. the basis of a large and prosperous 1961. The ultra-violet absorption of sea Japanese industry. But even the pearl water. J. Mar. Biol. Ass. U.K. 41(3): growers have their trials and tribula- 591-597. tions, for now and then the 'red tide' appears--an onslaught of red micro- Shows light absorption of micropore

organisms, so small that 25,000 of them (IfJ. ) filtered sea water at several wave have plenty of room to circulate in a lengths and suggests differences be- single cubic centinneter of sea water. tween water from different localities But they color the water red- -and they are due to organic matter. cause millions of dollars worth of ARNOLD, EDGAR L., JR. damage each year to the pearl-oyster 1958. Gulf of Mexico plankton beds. investiga- tions: 1951-53. U.S. Fish Wildl. Serv., "Japanese chemists have found that Spec. Sci. Rep. Fish. 269, vi + 53 they can halt the ravages of the 'red p. tide' by diffusing certain chemicals Presents the volumes of plankton, through the oyster bed. A solution of fish eggs, and fish larvae captured by ferric chloride does the trick; so does four types of collecting gear in 449 tows chlorine gas. Motor boats have been from the research vessel Alaska in the rigged up with apparatus to spread these Gulf of Mexico from March 1951 to July chennicals through the infested areas 1953. by discharging a stream of them fronn BALLANTINE, DOROTHY, and B. C. ABBOTT. the stern of the boat in such a way that 1957. Toxic marine flagellates; their oc- they are churned into the water by the currence and physiological propeller." effects on animals. J. Gen. Microbiol. 16(1):274- 281. ANONYMOUS. This paper includes a map showing 1953. Factors in "red tide" outbreak of "the world-wide distribution of 'red 1952. Progr. Fish-Cult. I5(3):128. tides' where mortalities have been shown to occur. ... 274.] A two-paragraph note on progress in [p. research during 1952.

20 "Blooms occur only when a number of Washington, and an analysis of diatona parameters are simultaneously suit- oil. Econ. Geol. 22(4):356-368. able " [p. 276.] Bioassays were conducted with toxin "The epidemics occur: from Gynnnodinium veneficum from the 1. Towards the end of the rainy Plymouth area. season in April and May. "Experiments have been carried out 2. After a heavy rainstorm. with a wide range of animals, and all 3. When the rains are followed by except polychaetes are affected to a gentle westerly winds. greater or lesser extent. In particular 4. And they reach a maximum when small fish, mainly gobies, have been the rain is followed by clear weather used as test animals; these die within and bright sunshine." [p. 358.] 10 min. in toxic cultures. Death in fish In 1925 thousands of tons of the dia-

seems to be due to some form of respira- tonns ( Aulacodiscus kittoni Arnott)were tory failure. The action of the toxin is washed ashore, covering the beach with certainly on the nervous system. The a mass 4 to 6 inches thick. first symptoms are a connplete loss of balance, combined with an intense BEIN, SELWYN JACK. change of colour pattern. This is fol- 1954. A study of certain chromogenic bac- lowed by a period of violent gasping, and teria isolated from "red tide" water with internnittent unco-ordinated bursts of a description of a new species. Bull. movement. During the quiescent inter- Mar. Sci.GulfCarib.4(2):110-119. vals, which become progressively longer, there is no response to sensory The author described a chromogenic stim\ilation. At the point of death there bacterium, which he named Flavobac-

is no sign of haemolysis in the blood, terium piscicida . This bacterium was the heart is beating, and excised muscles isolated from colored water with a respond to direct stimulation.... yellowish cast. He cultured the bacteria [p. 277.] and performed several experinnents in "... The toxin acts by depolarizing which fish died when held in 4-gallon excitable membranes, rendering them tanks to which a half-liter of a 24- hour inexcitable. Both nerves and muscles culture of the bacterium was added. He are affected, but in the intact animal suggested that since this bacteriunn is death occurs because of interference capable of killing fish under laboratory withtransnnission inthe nervous system. conditions it possibly had a role in the The effect is reversible in isolated mortality from "Red Tide" outbreaks. tissues, but the whole animal cannot dispose of the toxin. ..." [p. 278.] BEIN, SELWYN JACK. The authors conclude that "in the case 1955. Red tide bacterial studies. Univ. Mi-

of the G. veneficum toxin the effect is ami, Mar. Lab., Spec. Serv. Bull. 10: 1 -2. due to membrane depolarization rather than to a curare-like inhibition of re- "The conclusions drawn from these sponse to acetylcholine." [p. 280,] experiments are as follows: The fish are killed by a water soluble, toxic ma- BARKER, H. A. terial, produced by the growth of the 1935. The culture and physiology of the bacteria. The toxin is thermolabile, marine dinoflagellates. Arch. Mikro- that is, unstable at high temperatures, biol. 6(2):157-181. [Cited from Ryther, but extremely poisonous under natural 1955.] climatic conditions. Given the correct conditions these organisnns appear cap- Ryther said, "Barker (1935), who is able of 'blooming' and causing mass one of the pioneers in developing suc- mortality of marine organisms in na- cessful culture methods for dinoflagel- ture." lates, observed optimal tennperatures [p. 2.] for growth of some 14 species between 18° and 25° C." BEIN, SELWYN JACK.. 1957. The relationship of total phosphorus BAUGHMAN, J. L. concentration in sea water to red tide 1947. The Florida red tide. Texas Game blooms. Bull. Mar. Sci, Gulf Carib. Fish 5(12):6, 20-21. 7(4):3l6-329. BECKING, L. B., C. F. TOLMAN, H. C. McMILLIN, JOHN FIELD, and TADAICHI The total phosphorus concentrations HASHIMOTO. from both published and unpublished 1927. Preliminary statement regarding the data were exannined to note concentra- diatom "epidemics" at Copalis Beach, tion before, during, and after red-tide

21 outbreaks. He used data from Ketchum the Malabar and Kanara coasts and and Keen (1948), Graham et al. (1954), "red water" phenomenon as their prob- Chew (1953), Lackey and Hynes (1955), able cause. Proc. Indian Acad. Sci. and data takenby the Marine Laboratory, B31:339-50. [Cited by Ryther, 1955.] not already published, from the vicinity of Gasparilla Island in 1954 and 1955. Describes a bloom of Noctiluca mil- Total phosphorus values showed great iaris occurring along the Malabar Coast variation that appeared to have no rela- in October 1948, after the southwest tion to red tide. For instance, a series and before the northeast monsoon, dur- of 98 observations at 28 stations around ing calm, hot weather, with a salinity of Gasparilla Island in the fall of 1952, about 35 p.p.t. during a period without red tide, aver- aged 14.74 ug.at./l., with a maximunn of BONNOT, PAUL, and J. B. PHILLIPS. 60 ug.at./l. These are higher values 1938. Red water, its cause and occurrences. than the few observations in sea water Calif. Fish Game 24(l):55-59. from red-tide bloonns analyzed by Ketchum and Keen (1948). "During the last three weeks in August The fact that some reports con- and the first two weeks in September, sistently gave very low phosphorus 1937, the ocean water from Pt. Sur to values, while others gave consistently Pt. Reyes was affected in varying degree high values, causes us to speculate on with red water. It was not distributed the difference in analytical techniques uniformly throughout the above region that may have been used. but appeared in streaks and patches. It The author stated, "In periods of Red was first noted in Monterey Bay about Tide outbreaks the total phosphorus con- August 11, appearing as a streak paral- tent of a bloom, including the organisms leling shore. This streak was about 100 and particulate phosphorus, is lower yards wide and commenced approxi- than the nnaximunn values recorded in mately 200 feet offshore. Later this was 'normal' years and, with the exception scattered over a wider area, apparently of Ketchum and Keen's values, is not by tide and wind. Other streaks and significantly different from the adjacent patches appeared farther offshore. areas with no Red Tide examined at the Fishernrien reported seeing this dis- same period. At many times and places colored water as far south as Pt. Sur examined in the absence of Red Tides and as far north as Pt. Reyes. Areas of or bloonns, Florida west coast waters red water were also reported in the San contained at least as much and often Pedro region by W. L. Scofield, so it more than the estimated order of mag- appears that other localities along the nitude of total phosphorus contained in Pacific Coast were affected at this time. a heavy plankton bloom. The total phos- The presence of red water in the Monte- phorus values of Lackey and Hynes rey region during the above period was (op. cit.) for Red Tide stations, which the nnost extensive and persistent plague were analyzed with the organisms in the recorded for this area. samples, are unusually low for anytype "The organisms responsible for the of plankton bloom and there is no obvious red water epidennic in the Monterey- explanation for these figures." [p. 327- San Francisco region during August and 328.] September, 1937, have been identified

The author concluded, ". . .It is pos- by H. W. Graham of the Carnegie In- sible that the threshold level of total stitution, Washington, D.C., as of the phosphorus necessary to support dense genus Gonyaulax, species G. catanella populations of this organism is lower or a very sinnilar form." [p. 577] than originally assunned. The fact that the waters under consideration are BRAARUD, TRYGVE. usually rich in this mineral has perhaps 1945. A phytoplankton survey of the polluted led to much of the confusion in dealing waters of inner Oslo Fjord. Hvalradets with this problem. It seems very prob- Skr. 28, 142 p. [Cited from Ryther, able that, insofar as phosphorus is con- 1955.] cerned, the areas of the west coast of Florida which have recorded Red Tides BRAARUD, TRYGVE. are, at all times, capable of supporting 1951. Salinity as an ecological factor in an outbreak. Since the total phosphorus marine phytoplankton. Physiol. Plana- content may no longer be considered a tarium 4:28-34. [Cited by Ryther, limiting factor here it has no value in 1955.] predicting outbreaks." [p. 328.] BRAARUD, T., and 1. PAPPAS. BHIMACHAR, B. S., and P. C. GEORGE. 1951. Experimental studies on the dino- 1950. Abrupt set-backs in the fisheries of flagellate Peridinium triquetrum (Ehrb.) 22 .

"The greatest outbreaks of red water Lebour. Avhandl. Norsk. Vid.-Akad., probably occur toward the end of a phyto- Oslo, Mat.-Naturv. KLasse (1951) 2:1- plankton season; in areas where upwell- 23. [Cited from Ryther, 1955.] ing occurs during part of the year only, red water usually develops toward the BRAARUD, T., and E. ROSSAVIK. end or directly after the period of up- 1951. Observations on the marine dinoflag- welling " 953.] ellate Prorocentrum micans Ehrenb, [p. in culture. Avhandl. Norsk. Vid.-Akad., BRUNEL, J., G. W. PRESCOTT, L. H. TIFFANY Oslo, Mat.-Naturv. Klasse (1951) 1:1- ET AL. 18. [Cited from Ryther, 1955.] 1950. The culturing of algae. Charles F. Kettering Found, p. I-IX, 1-114. [Non BRONGERSMA-SANDERS, MARGARETHA. vidi.] 1945. The annual near Walvis bibliography on Bay (South West Africa), and its signifi- Contains an extensive and sumnnarizes work cance for paleontology. Arch. Neerland culturing of algae Zool. 7:291-294. [Cited from Pomeroy on culturing. et al.. 1956.] BRYAN, ARTHUR H. Biol. Teacher Upwelling of water into photic zone is 1963. The red tide. Amer. Abstr. given as a source of nutrients. 25(l):53-54. [Cited from Biol. Non vidi.] BRONGERSMA-SANDERS, MARGARETHA.. and 1948. The importance of upwelling water to BUCK, JOHN D., SAMUEL P. MEYERS, vertebrate paleontology and oil geology. EINAR LEIFSON. pisci- Verhandl. Koninkls. Nederland. Akad. 1963. Pseudomonas ( ) Wetenschap., afd. Natuurk., Tweede cida Bein Comb. Nov. J. Bacteriol. Sectie 45(4):1-112. [Cited by Ryther, 86(5):1125-1126. 1955.] Discovered the organism to be polar peritrichous Blooms of Noctiluca sp. appeared to monotrichous rather than cause annual mass mortalities in Walvis as originally described by Bein (1954). Bay, South West Africa, in December. Culture filtrates were toxic to the top Mortalities were during periods of mini- minnow Gambusia sp. mum upwelling and the highest water temperature of the year. BURKHOLDER, PAUL R., and LILLIAN M. BURKHOLDER. MARGARETHA. BRONGERSMA-SANDERS, 1956. Vitamin Bj2 ^'^ suspended solids and 1957. Mass mortality in the sea. In Joel W. marsh muds collected along the coast of Hedgpeth (editor), Treatise on marine Georgia. Limnol. Oceanogr. 1(3):202- ecology and paleoecology, vol. 1, ch. 208. 29, p. 941-1010. Geol.Soc. Amer., Mem. 67. Appreciable amounts of vitamin B12 Mass mortality in sea caused by are carried on suspended particles in noxiousness of waterbloom [p. 951], river water, up to 6.4 micrograms per "Red water occurs in very fertile gram of solids. On particulate matter in parts of the sea, often during or after sea waters vitamin Bi2 varied from unusually warm weather; it develops in 0.0027 to 0.130 microgram per liter. succession after a great production of other organisnns. Products of decay of BURR, J. G. the latter might be one factor that sets 1945. Science tackles a mystery. Texas the bloom in motion. Great outbreaks Game Fish 3(9):4-5, 24-25. occur particularly in subtropical and tropical regions where the rate of over- "On June 27, 1935, ten years ago, the turn of organic matter is high; in fer- Texas coast was rocked by the nnost tile parts of high latitudes red water, at sensational event in its history, when least of catastrophic proportions, does millions of pounds of fish were destroyed not occur or is very rare. Plenty of in the Gulf of Mexico by some xanknown sunshine seems another requirennent for cause. This destruction of fish con- red-water outbreaks. The areas where tinued for a period of five weeks or red water occurs are somewhat renni- more as evidenced by dead fish which niscent of polluted waters; the dead fish lined the shore for 150 nniles, beginning will worsen the 'pollution'. Here it is along the shore of Padre Island, and important to note that pollution by extending nearly to the area of Free-

human action also favors red-water port. . .for two months prior to the fish

outbreaks. . . tragedy rainfall had been unprecedented. 23 and the four major rivers from the CARTER, H. J. Brazos to the Nueces River, in the 1858. Note on the red colouring matter of months of May and June, had dunnped the sea round the shores of the island 10,000,000-acre feet of water into the of Bombay. Ann, Mag, Natur. Hist,, Gulf." [p. 4.] 3d ser., l(4):258-262. BURSA, ADAM. He attributed the discolored water to 1963. Phytoplankton in coastal waters of the a form which he named Peridinium

Arctic Ocean at Point Barrow, Alaska. sanguineum . Arctic l6(4):239-262. CHEW, FRANK, Lists 73 species (one new) in 20 genera 1953. Results of hydrographic and chemical of dinoflagellates collected between mid- investigations in the region of the "Red Jvuie and mid-September of 1954. Maxi- Tide" bloom on the west coast of Florida mum temperature in August was 10.2'-* C. in November 1952. Bull, Mar, Sci, Gulf CABASSO, v., and H. ROUSSEL. Carib. 2(4):6l0-625. 1942. Essai de explication du phenomene dit He reported the "signs of another "Des eaux rouges" du lac de Tunis. bloom off Dry Tortugas on January 3, Arch. Inst. Pasteur, Tunis, 31 (3/4):203- 1953, and subsequent airplane observa- 211. tions disclosed suspicious areas, A CAHN, A. R. definite recurrence was detected off 1949. Pearl culture in Japan. Natur. Re- Smith Shoal on January 22, 1953. This,

sourc. Sect. Supreme Comnnander Allied too, was of brief duration. . . ." [p. 6IO,] Powers, Tokyo, Rep. 112, 91 p. Off Fort Myers red tide occurred in November 1952; 11 hydrographic sta- Describes the destruction of pearl tions were occupied (actually 12 sta- oysters by red tide. tions). CAHN, A. R. Each of the 12 stations was occupied 1950. Oyster culture in Japan. Fish Wildl. once during a period of 3 days. The Serv., Fish. Leafl. 383,80 p. depths were 7 to 13 m. and observations "The 'akashio' or red tide is an of salinity, temperature, and total phos- aquatic phenomenon that occurs at ir- phorus were taken at surface, bottom, regular intervals and with variable in- and middepth. The author attempted by tensities and results. It nnay be highly nneans of hydrographic sections to postu- destructive not only to edible oysters but late "river water" and "Gulf water" in also to the pearl oysters (Cahn, 1949) the snnall near- shore area involved; and to other forms of . In however, the few data, the slight dif- recent years the microorganisms which ference in salinity, and the use of cause the red tide in Japan have been straight line interpolation of values shown to be various dinoflagellates such where there is no consistent pattern,

as Gonyaulax , Gymnodinium , Peridi- all make the exercise appear tenuous,

nium , Ceratium , and other forntis of lesser importance, [p. 60,] CHEW, FRANK. 1955a, On the offshore circulation and a convergence mechanism in the red tide region off the west coast of Florida. "Red tides occur mostly in the spring Trans. Amer. Geophys, Un, 36(6):963- and autunan where the ocean current is 974; also Tech. Rep. 55-5, Fla, State sluggish, usually during continuously Bd. Conserv., 27 p. fine, quiet weather. . . . Many red tides do little or no damage to oysters, while others result in a mortality approach- In this articlethe author discussed a ing or reaching 100 percent. ... Experi- mechanism to concentrate floating par- ments using copper sulfate are incon- ticles. He quoted from several obser- clusive." [p. 60.] vations concerning floating dead fish, and then reached the conclusion that, CANOVA, P. ANDREW "The large horizontal dimensions of 1885. Life and adventures in south Florida. these patches of dead fish preclude the South Siin Publ. House, Palatka, Fla. phenonnenon from being oversized wind- 136 [Cited from Lackey and Hynes, p. rows or resulting from causes related 1955.] to the wind [Stommel, 1951].... Canova reportedly mentioned a "green In the next paragraph he stated, "The tide" in 1885 that might have been as observed outbreaks of Red Tide at intense as later outbreaks butless irri- beaches generally show a sequence of ." tating to the then small tourist industry. south to north movennent. , . Recent 24 outbreaks have not shown such tendency. theory of red tide origin and water move - The author then said, "The renaaining ment. portion of this paper discusses the off- CHEW, FRANK. shore circulation in an attempt to find 1955b. Red tide and the fluctuation of con- the northward and the mechanics of servative concentrations at an (italics ours), shoreward movement," mouth. Bull. Mar. Sci. Gulf Carib. 5(4): [p. 963.] 321-330. In advancing his thesis the author placed heavy weight on the few phos- This paper was based on salinities phorus samples analyzed by Ketchum taken in a row of eight stations on May and Keen (1948). Actually, the values 28, 1954, over one tidal cycle of 14 which Ketchum and Keen observed were hours. The paper attempted to showthat not as exceptional as comment at the if red tide requires aparticvilar salinity, tinne would have us believe. Their values such as that found at the mouth of an for total phosphorus in red-tide water estuary, this salinity may be found best have often been exceeded by samples at times of greatest mixing. This paper from nonred-tide water collected by the stressed salinity as the cause of red Bureau of Commercial Fisheries. Ex- tide. Other explanations, however, were ceptionally high phosphorus values often also given. For instance he stated, "It occur in nonfiltered samples because of has been suggested that observed Red particulate matter. This refutes their Tide outbreaks have tended to fall near statement, quoted in Galtsoff (1948, periods of spring tides. Since from equa- p. 21)," ... These values are from 5 to tion (11) one sees that the transition 10 times as high as those ever en- zone near the estuary mouth is weak countered in uncontaminated oceanic during these periods, one is led to the water. ..." conclusion that an effect of spring tide In advancing his theory of the offshore on Red Tide outbreaks is due to other derivation of phosphorus-rich water. causes, such as stirring of the estuary

Chew stated, ". . . The shoreward ten- bottom, than the presence of a strong, dency during a Red Tide bloom was sharp transition zone at the estuary noted earlier by Chew [1953] who con- mouth." [p. 329.] cluded that high total phosphorus waters CHEW, FRANK. were advected toward shore from the 1955c. The summer circulation of the open Gulf; the of and work Ketchum Florida west coast offshore water as Keen [1948] is also of significance in deduced from the pattern of thermo- this respect, since their highest ob- cline depths and a non-geostrophic equa- served total phosphorus was found at tion of motion. Rep. to Fla. State Bd. their farthest offshore stations " Conserv., Tech. Rep. 55-12, 6 p. [p. 972.] A completely opposite picture would be obtained from the data from This paper advances a theory of the 16-month survey of phosphorus by water circulation based chiefly on BT the Bureau of Commercial Fisheries readings from one August cruise of the

(Graham et al., 1954). vessel Alaska . The tinne period was In the discussion of water movements not mentioned but from the markings on Chew did not separate floating dead fish his figure 2 it appears to be about 1 from red tide. Actually, they are not the week. This paper nnarks a step toward same and cannot be expected to act in advancement of Chew's theory that the the same way. cause of red tide must be sought else- In his summary Chew stated that where, such as the Apalachicola River. "... The mortality generally moves CHEW, FRANK. northward and shoreward. ..." 973.] [p. 1956. A tentative method for the prediction Regarding this conclusion he cited of the Florida Red Tide outbreaks. Gunter et al. (1948) and one issue of a Bull. Mar. Sci. Gulf Carib. 6(4):292-304. St. Petersburg newspaper. Chew also made a second conclusion In this paper the hypothesis is ad- in which he said, "During periods of vanced that red-tide outbreaks are Red Tide mortality, it is generally be- caused by the mixing of estuarine waters lieved that there is a large band of dead carrying a terrigenous nutrient supply, fish some tens of miles offshore. The with offshore waters. The Apalachicola horizontal dimensions of the band are River was taken by Chew as the source several miles wide and 20 to 30 mi. of the estuarine water. He said it is long...." [p. 973.] See, for example, "seeded" by estuarine waters from Nicholson (1954). This band of dead fish Tampa Bay and Charlotte Harbor which appears to be a necessary part of his contain the causative organism. His Red

25 " Red tide research. --Data collected Index Number was synthesized Tide the November 1952 red tide out- from Apalachicola River runoff data during break, which also killed millions of and air temperatures for several locali- fishes off Florida in 1946 and ties. The Index appears to have been 1947 and has been ascribed to an excess derived by working backward from ef- of the dinoflagellate Gymnodinium bre- fect to cause, and no measure was given vis, show that Caloosahatchee River of its reliability. One of the chief dif- effluents are important agents in such ficulties in deciding on causal factors and that organic content and has been the lack of good data on abun- blooms, physical attributes cause such activity. dance of red-tide organisms. tank work indicates a His unsubstantiated theory requires Experimental growth of dinoflagellates as well chunks of Apalachicola "estuarine mass that organisms, requires high light water" move intact from Apalachicola as other intensity, vitamin B12 and sulfides. to the vicinity of Boca Grande Pass and of tidal streams, there acquire red-tide organisms. "An examination marshes and produced Gym- CHIDAMBARAM, K., and M. D. ANNY. nodinium brevis in Barfield Bay, south plank- 1944. Note on the swarming of the Florida. Field culture of this species tonic algae, Trichodesmium erythraeum was unsuccessful. Water from Lake in the Pamban area and its effect on the Okeechobee was heavily loaded with dis- fauna. Curr.Sci., Bangalore, 13(10):263. solved organic materials. and CLASSEN, TH. "Daily collections and physical were made of 1930. Periodisches Fischsterben in Walvis chemical observations Texas, la- Bay, South West Afrika. Palaeobiologica dinoflagellates in Galveston, used in 3:1-13. goons. Gymnodinium splendens , experiments as nearest to Gymnodinium CLEMENS, W. A. brevis, was found. 1935. Red "water-bloom" in British Colum- bia waters. Nature 135(3412):473.

"In Nature of September 22, 1934, " Microbiology .- -Light levels above there is a communication from Mr. T. those generally reported for algal cul- of John Hart describing the occurrence ture have been found necessary for cul- causedby Mesodin- a red 'water-bloom' ture of some dinoflagellates indigenous African seas. It may be ium in South to the Gulf coast. A ten-fold increase is an occurrence of of interest to record now in use. Large-scale tank studies blood- red water at Nanaimo, British have proved valuable adjuncts to stand- Columbia, during the week of April 28, ard tube and dish cultures. 1933. The water in a channel immedi- " Experimental ecology .-- These ex- ately north of the harbour was coloured periments have been confined to a study in great patches. Examina- crimson red of red tide. Large tanks have been set of the water revealed tion of a sample up for use as a 'stepping stone' from of a ciliate, identified by a pure culture test tube to open sea. Findings on light as Mesodinium rub- Mr. G. H. Wailes level and value of certain inorganic rum , Lohmann. nutrients result from this approach. in Lady- "About this time oysters These are important in analyzing inter- south of smith Harbour, fifteen miles species on relationships. This work reported to contain red Nanaimo, were provides a basis for advanced studies 'worms.' Investigation disclosed the of larval shrimp and fish behavior, fact that the crystalline styles were survival and growth." [p. 21.] coated with a red colouring matter, evidently as a result of feeding upon COLLIER, ALBERT. and zirconium in bloom of Mesodinium . Examination of the styles 1953b. Titanium of local clams showed a similar condi- Gymnodinium brevis Davis. Science tion. 118(3064):329. of this 'bloom' of "The appearance Samples of water were analyzed from: Mesodinium followed a period of a 1. A dense bloom of G. brevis couple of weeks of bright, sunny, calm 2. Lake Okeechobee weather. No discoloration was observed 3. Central Gulf of Mexico in 1934." 4. A tidal lagoon at Galveston, Tex. COLLIER, ALBERT W. Titanium and zirconiunn were suf- 1953a. Gulf Fishery Investigations. In An- ficiently abundant in sample 1 to exceed nual report for fiscal year 1953, Branch the sensitivity of the analytical method. of Fishery Biology, Fish Wildl. Serv., No traces of either could be detected p, 20-21. in the other four samples.

26 COLLIER, ALBERT W. "Laboratory experiments have fur- 1954. Gulf Fishery Investigations. &i Annual ther confirnaed the theory that decaying report for fiscal year 1954, Branch of fish bodies reinforce the bloonns." Fishery Biology, Fish Wildl. Serv., [p. 24-25.] p. 23-25. COLLIER, ALBERT W. " Red Tide .- -When the September 1953 1955. Gulf Fishery Investigations, bi An- Red Tide outbreak occurred the micro- nual report for fiscal year 1955, Branch organisna Gynnnodinium brevis was iso- of Fishery Biology, Fish Wildl. Serv., lated and a medium evolved in which it p. 29-32. could be grown as a unialgal culture (not bacteria free). In a series of ex- "Support for the theory that residues periments, tube cultures were enlarged of dead fish con-ipound the division rate to 10-liter tank cultures which culmi- of G. brevis comes from an explosive nated in developnnent of small-scale bloom in the laboratory which was blooms which were toxic to fish. The created by adding juices from fish success with Gymnodinium brevis was killed experimentally by a relatively made possible by success with cultures low concentration of G. brevis . The of a related species, Gymnodinium initial concentration was about 3,000,000 splendens . Cultures of the latter tested individuals per quart; with the addition for toxicity on fish (Membras and Mol- of juices they tripled in nunaber over- lienesia ) were nontoxic at concentra- night. tions of 3,000,000 cells per liter. Gymnodinium brevis was toxic at

1 ,300,000 cells per liter, a concentration " which does not cause a pronounced water Red tide control . --K control is pos- discoloration in the 5- gallon pyrex con- sible, it will come after development of tainer where the culture is maintained. techniques for predicting conditions In the deeper waters of the Florida which are conducive to a red tide out- littoral regions, however, this concen- break and for economical distribution tration would cause the 'milky green' of chemical or physical agents lethal to

which often characterizes areas where G . brevis . Predictions are necessaryto fish are killed. make the latter possible. "These mass laboratory cultures are set up so cells can be harvested daily and the volume of total culture re- moved made up by adding fresh nriedium. "A search for lethal agents has re- Daily and biweekly sampling has been vealed that metallic copper and copper made for pH, carbon dioxide, nitrate salts are most toxic. Copper sulphate nitrogen, phosphate phosphorus, car- at approximately 0.12 p.p.m. (copper bohydrate, tyrosine, ammonia, and sul- equivalent 0.05 p.p.m.) in sea water is

fide in these cultures. Metabolic studies lethal to G. brevis . A small piece of onmass cultures are used in interpreting copper metal will make a pint of sea field data and will assist in diagnosing water lethal to G. brevis in a few field conditions which can lead to a seconds. An experiment in which fine Red Tide. copper sulphate was spread over the "Field surveys which began with the sea surface with crop-dusting planes September 1953 outbreak have shown was successful in eliminating the organ- Florida waters have never been entirely isms for approximately one fourth of a

free of Gymnodiniunn brevis . Although square mile." [p. 31.] suitable hydrographic conditions have been present to some degree through COLLIER, ALBERT. the whole period, it is unknown whether 1958a. Gulf of Mexico physical and chemi- continued presence of G, brevis is be- cal data from Alaska cruises. U.S. cause of these conditions or because of Fish Wildl. Serv., Spec. Sci. Rep. Fish. improved techniques for locating the 249, vi + 417 p. organism when it is not in bloom. Data from laboratory and field studies sup- This reports the original hydro- port the tentative conclusion that op- graphic data collected from the vessel timum salinity for a bloom of this Alaska during the period March 1951 to organism lies between 32 and 34 parts July 1953. per thousand. "Analysis of clinnatic factors has COLLIER, ALBERT. confirmed the theory that rainfall dis- 1958b. Some biochemical aspects of red tribution is a prime factor in initiating tides and related oceanographic prob- fish kills. lems. Limnol. Oceanogr. 3(l):33-39. 27 The frequency distributions of pollutant entering the bayou It sannple values for carbohydrates, tyro- seems logical to assume that these sine, and organic phosphorus are given, extraordinarily high values of BOD and and the author stated that "there was a consequent anaerobic conditions in the tendency toward the contagious distri- water were attributable to the Gonyau- butions." lax and were important contributing Plotting shows that the carbohydrate factors in the mass mortality of fish in values plotted logarithmically form a Offatts Bayou." [p. 362-363.] linear curve. Organic phosphorus defi- nitely does not, and the tyrosine values CONOVER, SHIRLEY A. MacMILLAN. do not, show a good linear fit. If, as 1954. Observations on the structure of red nnany authors contend, plankton is dis- tides in New Haven Harbor, Connecti- tributed in one of the logarithmic series, cut. J. Mar. Res. 13(1):145-155. it would not be surprising if carbo- hydrates, if derived from plankton, also "...High concentrations of red tide fell into the two-parameter logarithmic organisms, here two species of Gon-

series. iaulax , were found to be associated with It was suggested that lethality of sea stable water masses of inner harbor water from different localities may be origin. The configuration of the harbor caused by the interplay of lethal con- permits the developnnent and retention centrations of copper and H2S; also of water masses; this tendency is rein- other "natural organic compounds" may forced by large volumes of nutrient- act both as chelators and growth-pro- rich fresh water entering the inner moting substances. harbor, by certain wind patterns, and by It was stated that W. B. Wilson iso- high radiation values. Adequate time, lated and cultured G. breve from Gal- illumination, nutrients, and favorable veston waters and again from waters of temperatures are essential for red tide "South Texas and Mexican east coasts." development...." [p. 145.] The author stated that heavy blooms of red tide "may augnnent themselves CORNMAN, IVOR. with both inorganic and organic ma- 1947. Retardation of Arbacia egg cleavage terials produced by the decomposition by dinoflagellate-contaminated sea products of dead fish." water (red tide). [Abstract.] Biol. Bull. (Woods Hole) 93(2):205. COLLINGSWOOD, C. 1868. Observations on the microscopic alga Samples of red tide sea water sup- which causes the discoloration of the plied by Paul S. Galtsoff were used at sea in various parts of the world. Trans. the Marine Biological Laboratory in Roy. Microscop. Soc, n.s., 16:85-92. Woods Hole. A sample of untreated decomposing sea water "taken fronn an area stained red by Gymnodinium when CONNELL, CECIL H., and JOY BARNES diluted 1:10" increased cleavage time CROSS. of the eggs of Arbacia by 1 hour (100 1950. Mass mortality of fishassociated with percent), "if added 10 minutes after the protozoan Gonyaulax in the Gulf of fertilization." However, "when most of Mexico. Science 112(2909):359-363. the H2S was pulled off with a vacuum pump, the delay at 1:10 was only 3 Contains a report of fish kills in minutes" and even at 1:5 only 15 min- Offatts Bayou associated with Gonyau- utes. lax catenella (?) during the summer of 1949. COVELL, W. P., and W. F. WHEDON. The authors indicated that: 1937. Effects of the paralytic shellfish poi- 1) Growth of Gonyaulax was increased son on nerve cells. Arch. Pathol. 24(4): by small amounts of sewer pollution. 412-418. [Non vidi.] As long as the organisms remained 2) COWEY, C. B. near the surface, the dissolved- oxygen 1956. A preliminary investigation of the content of the water was high. However, variation of Vitamin B12 in oceanic and when the organisms submerged, the coastal waters. J. Mar. Biol. Ass. oxygen content rapidly approached zero. U.K. 35(3):609-620. [Cited from Mc- Only seven samples were taken in all. Laughlin, 1958,] 3) "The values of biochemical oxygen demand (BOD) for the samples asso- GRANGE, JOHNIE H. ciated with Gonyaulax were extraordi- 1963. The effects of copper sulfate on nary and far in excess of what might be Microcystis and in ponds. explained by sewage or any other organic Progr. Fish-Cult. 25(4):198-202.

28 Copper sulphate (CUSO4.5H2O) at a Gymnodinium brevis amounted to 99.28 concentration of 0.05 p. p.m. killed Mi- per cent and 98.99 per cent of the total crocystis without harming copepods or organisms in the water in two uncon- cladocerans. centrated samples of sea water." [p. 83.] DARESTE, CAMILLE. 1 855. Memoire sur les animalcules et autres DAVIS, CHARLES C, and ROBERT H. corps organises qui donnent a la mer WILLIAMS. une couleur rouge. Ann. Sci. Nat., Zool. 1950. Brackish water plankton of mangrove 3:179-239. [Cited from Martin and areas in southern Florida. Ecology Nelson, 1929.] 31(4):519-531.

Gives numerous references to early Plankton samples were writers and navigators concerning red collected dur- ing the first half of 1948 (plus four tide, samples in June 1947) from 28 locali- ties in the mangrove area of the Ever- DAVIS. CHARLES C. glades, between Everglades City and 1948. Gymnodinium brevis sp. nov.,acause . Salinities are given on of discolored water and animal mor- dates of collection. tality in the Gulf of Mexico. Bot. Gaz. 109(3):358-360. [Contr. 17, Univ. Mi- De SOUSA e SILVA. ami, Mar. Lab.] 1953. "Red water" par Exuviella baltica "In April, 1947, there were further Lohm, Com simultanea mortalidade de reports of fish mortality in the Gulf of piexes nas aguas litorais de Angola.

Mexico off the Florida Keys. . . . Trab. MissSo Biol. Marit. 4, Lisboa. "... Off Key West (near Content Keys) [Non vidi.] fish mortality was occurring in a situa- Analyzed red-tide plankton samples tion in which the count of Gymnodinium from Luanda (see Niimann, 1957). was 420,000 cells per liter. . . Further- more, the mortality of fish and other DENISON, W. animals occurred sporadically over a 1862. On the death of fishes during the period of 9 nnonths from November, monsoon off the coast of India. Quart. 1946, to July, 1947. . . ." [p. 358,] J. Geol. Soc, Lond. 18:453. Gymnodinium brevis is described as a new species, with an illustration. The DRAGOVICH, ALEXANDER. chromatophores are described as 1958. Hydrography related to red tide. In yellow- green in color in both living and preserved specimens. Annual report of the Gulf Fishery In- vestigations for the year ending June 30, 1958, U.S. Fish Wildl. Serv., 70-75. DAVIS, CHARLES C. p. 1950. Observations of plankton taken in Copper, total phosphate-phosphorus, marine waters of Florida in 1947 and inorganic phosphate-phosphorus, and 1948. J, Fla, Acad. Sci. 12(2):67- 1 03. nitrate- nitrites in the coastal water of west Florida are subjects of this report. Lists plankton species taken in Flor- ida waters (chiefly west coast) during DRAGOVICH, ALEXANDER. 1947. About half the samples were taken 1959. Environmental hydrology in relation with a fine net (No, 20) and several of to red tide, in Galveston Biological them contained large numbers of Laboratory fishery research for the Gymnodinium breve. year ending June 30, 1959, p. 80-85. U.S. Fish Wildl. Serv., Circ. 62. DAVIS, CHARLES C. 1955. The marine and fresh-water plank- "One of our objectives is to deter- ton. Mich. State Univ. Press, 562 p. mine whether the outflow of the Hills- boro, Alafia, Little Manatee and Mana- In early May 1948 the water of Middle tee Rivers influences the phosphorus Lake, Dade County, was green and content of Tampa, Bay and adjacent opaque owing to the enormous abundance neritic waters The water of upper

of Gonyaulax , The water of certain areas Tampa Bay (subarea 2) contained nnore of Seven Palm Lake showed similar phosphorus than any of the rivers, ex- patches. cept the Alafia. These data indicate that "Unpublished data in the possession phosphorus may, on occasions, accumu- of the author show that during the 'red late in this portion of Tampa Bay. tide' on the west coast of Florida, de- scribed by Gunter et al. (op. cit.),

29 "In March, a month of heavy rainfall, DRAGOVICH, ALEXANDER. a drastic increase in phosphorus content 1963. Hydrology and plankton of coastal was noted in all areas except at stations waters at Naples, Florida. Quart. J. located 20-40 miles offshore. In April Fla. Acad. Sci. 26(l):22-47. and May, the precipitation continued to In this report on plankton at Naples be heavy and in May a nominal increase from March 1956 to August 1957 the content in the areas 20 to of phosphorus following observations on occurrence ." 40 miles offshore was observed. . , of Gymnodinium breve in certain tem- [p. 82.] perature and salinity ranges are of DRAGOVICH, ALEXANDER. interest: 1960a. Copper in Tampa Bay and adjacent Temperature ranges ('''C.) neritic and river waters, ij Galveston 17.0-23.5 23.6-28.4 28.5-31.9 Biological Laboratory fishery research for the year ending June 30, 1960, Total p. 65. U.S. Fish Wildl. Serv., Circ. 92. numbers of samples 126 120 125 Information in this progress report Samples published in Dragovich and May (1963). with G. breve 3 9 5 DRAGOVICH, ALEXANDER. 1960b. Hydrology of Tampa Bay and adjacent Salinity ranges (p.p.t.) waters. In Galveston Biological Labora- 32.4-36.0 36.1-36.9 37.0-38.6 tory fishery research for the year end- ing June 30, 1960, p. 48-51. U.S. Fish Total Wildl. Serv., Circ. 92. numbers of samples 126 121 115 "Connparison of the water tempera- Samples tures with incidence G. breve indi- of writh G. cates that a relation to temperature may breve 9 4 1 exist, especially during the cold periods. During the winter [59-60] the tennpera- ture of water 20-40 miles offshore was DRAGOVICH, ALEXANDER, JOHN H. FINU- higher than that of inshore waters. This CANE, and BILLIE Z. MAY. may be important in the survival of G. 1961. Counts of red tide organisms, Gymno-

breve offshore. . . ." [p. 48.] dinium breve , and associated oceano- It was suggested that "the size of a graphic data from Florida west coast, G. breve population may be important 1957-59. U.S. Fish Wildl. Serv., Spec. for its survival in Tampa Bay during Sci. Rep. Fish. 369, iii + 175 p. the onset of a rainy season when a drastic reduction of salinity occurs," This report continues the presenta- [p. 49.] tion of original data (see Finucane and It was also suggested that it is pos- Dragovich, 1959) on red tide and asso- sible for surface waters with a slightly ciated ecological conditions. It covers reduced salinity to exist at 30 to 40 the period from July 1957 through miles offshore. December 1959 and the area from The author said that the greatest Anclote Keys to Cape Sable. amount of phosphorus is brought into DRAGOVICH, ALEXANDER, JOHN H. FINU- Tampa Bay by the Alafia River, and the CANE, JOHN A. KELLY, JR., and BILLIE greatest proportion of phosphorus is Z. MAY. introduced at the beginning of the rainy 1963. Counts of red-tide organisms, season. Gymnodinium breve , and associated "... No relationship was observed oceanographic data from Florida west between the incidence of G. breve and coast, 1960-61. U.S. Fish Wildl, Serv., nitrate-nitrites." [p. 51.] Spec. Sci. Rep. Fish. 455, iii + 40 p. DRAGOVICH, ALEXANDER. This paper continues the presentation 1961. Relative abundance of plankton off of original data on red tide and asso- Naples, Florida, and associated hydro- ciated oceanographic data from the west graphic data, 1956-57. U.S. Fish Wildl. coast of Florida (see Finucane and Serv., Spec. Sci. Rep. Fish. 372, Dragovich, 1959, and Dragovich, Finu- iii + 41 p. cane, and May, 1961). This report, how- Original data from Naples, Fla., for ever, includes observations on am- the period March 1956 through August monia, total and inorganic nitrogen, 1957. silicon, calcium, and alkalinity.

30 DRAGOVICH, ALEXANDER, and JOHN A. district, than in the Hillsborough River. KELLY, JR. All of the rivers entering Tampa Bay 1964. A collection of data in reference to have higher phosphorus than the Peace red tide outbreaks during 1963. 2. Pre- and Caloosahatchee Rivers. liminary observations on phytoplankton "... Results of this investigation have and hydrology in Tampa Bay and the shown that the average concentration of immediately adjacent offshore waters. copper for all rivers combined is well ." Fla. Bd. Conserv., Mar. Lab., p. 4-22. below the toxic levels for G. breve . . . [p. 175.] From January to May, 1963, gives temperature, salinity, oxygen, inorganic DROOP, M. R. phosphate-phosphorus, chlorophyll A, 1954. A note on the isolation of small and monthly precipitation. Eight sta- marine algae and flagellates for pure tions were occupied monthly. Blooms cultures. J. Mar. Biol. Ass. U.K. 33(2): in April of Gymnodinium breve coin- 511-514.

cided with blooms of Ceratium furca ,

Prorocentrum micans , Gonyaulax die- EICHER, GEORGE. in aquatic weed control. gensis (?) Rhizosolenia stolterfothi , and 1947. Aniline dye Tintinnidae. The red tide coincided with J. Wildl, Manage. 1 1(3):1 93- 197. [See an intrusion into Tampa Bay of offshore section of this report on control water at a period of exceptionally low methods.] rainfall. ELDRED, BONNIE, KAREN STEINDINGER, DRAGOVICH, ALEXANDER, and BILLIE Z. and JEAN WILLIAMS. MAY. 1964. A collection of data in reference to 1961. Hydrology of Tampa Bay and adjacent red tide outbreaks during 1963. 3. Pre- waters. In Galveston Biological Labora- liminary studies of the relation of tory fishery research for the year end- Gymnodinium breve counts to shell- ing June 30, 1961, p. 65-67. U.S. Fish fish toxicity. Fla. Bd. Conserv., Mar. Wildl. Serv., Circ, 129. Lab., p. 23-52.

"... The distribution of G. breve Toxicity of oysters rose when red seems to be restricted by temperature tide occurred in April to December, minima rather than maxinna. G. breve 1963 and decreased rapidly when red was absent at temperatures below 14.2° tide diminished. C, which occurred in December and present at 31.3° C. in July. ELDRED, BONNIE, VIOLET STEWART, "G. breve was absent from Tampa LARRY GILLESPIE, JEAN WILLIAMS, and Bay during the months with lowest KAREN STEINDINGER. temperatures. The organism's simul- 1964. A collection of data in reference to taneous presence in the farther offshore red tide outbreaks during 1963. 4. Pre- waters continues to suggest that the liminary studies of vitamin Bj2. carbo- warmer neritic waters maybe important hydrate, and phytoplankton in sea water. Mar. Lab., to the winter survival of G. breve . Fla. Bd. Conserv., "... During most of the year precipi- p. 53-96. tation was below normal or near to the Extensive tables for May to Septem- Extremely heavy long-term means. ber, 1963, giving for several inshore rainfalls were associated with hurri- and river mouth stations, vitamin B 12 cane 'Brenda' (July 28 and 29). Accord- (assayed with Cyclotella nana ), carbo- rainfall, hurricane ing to the reported hydrates (method of Lewis and Rake- (September 9-11), which swept 'Donna' straw, 1955), phytoplankton counts by drain- through the general Tampa Bay large groups, salinity, and temperature, age basin, was not a particularly wet

hurricane. . . ." [p. 65.] ENDLICH, F. M. 1882. An analysis of water destructive to DRAGOVICH, ALEXANDER, and BILLIE Z. fish in the Gulf of Mexico. Proc. U.S. MAY. Nat. Mus. 4:124. Hydrological characteristics of 1962. He reported upon two samples of sea Bay tributaries. U.S. Fish Wildl. Tampa water, one in which fish died and another Fish. Bull. 62:iv 163-176. Serv., + p. in which they did not. He concluded that "I find that the water A [in which fish Higher concentrations of total and died] contains a large quantity of Algae inorganic phosphorus occurred in the and Infusoria. . . . Alafia, Little Manatee, and Manatee Rivers, which flow through a phosphatic

31 "In my estimation the death of fish Their conclusion that the outbreaks was caused by the more or less para- "seem to move from south to north" sitic alage, which are found in large does not appear to be justified by the quantities in water A, but do not occur evidence presented. The authors appar- at all in water B." ently recognized this weakness, stating that "Further data are required to give EWING, GIFFORD. complete support to Z." [p. 1,] 1950. Slicks, surface films and internal In figure 3 they presented a histogram waves. J. Mar. Res. 9(3):l6l- 187. of the number of reported occurrences by month from 1878 to 1953, inclusive. Explains the physical basis and bio- The data give an incorrect innpression, logical significance of most slicks. because the authors apparently listed the same nnonths as many times as they FEINSTEIN, ANITA. ran across a report. The difference 1956. Correlations of various ambient phe- between their histogram and the tabula- nomena with red tide outbreaks on the tion we get for the same data for 1878- Florida west coast. Bull. Mar. Sci. 1954 is as follows: Gulf Carib. 6(3):209-232.

Jan. Feb . Mar. Apr. May June The author calculated linear correla- tions between red tide outbreaks and Report rainfall, tropical disturbances, and Our tabulation

stream discharge. In table 1 the author listed all years with red-tide outbreaks July Aug. Sept. Oct. Nov. Dec. and ranked their intensity on a scale from to 5 from 1844 to 1954. The Report data in the table are unavoidably sub- jective. The author, for instance, re- corded a light outbreak in 1949 based on only one report of 3,000 dead fish (cause of death not certain) on Anna

Maria Beach. Although 1 887 is not listed in the table, the text mentions 1887 data from Harper (1927). For two of the years (1931 and 1936) the author gave as authority a personal communication (un- identified writer) to a member of the laboratory staff (also iinidentified). This lack of adequate documentation, although not the fault of the author, is unfortu- nate. The difficulty in measuring the abun- dance of the red tide organisms by re- ports of fish kills is probably so great that correlations with other phenomena are necessarily hard to detect.

FEINSTEIN, ANITA, A. RUSSEL CEURVELS, ROBERT F. HUTTON, and EDWARD SNOEK, 1955. Red tide outbreaks off the Florida west coast. Univ. Miami, Mar. Lab., Rep. 55-15 to Fla. State Bd. Conserv., p. 1-44.

The authors gave a chronological report on red-tide occurrence derived from many sources from 1844 to Janu- ary 1955. Admitting that the data are subjective, the authors attempted to list the intensity of red-tide occurrences in

categories of magnitude from 1 to 5 by calendar years.

In figure 1 they showed the occur- rences by month from December 1953 to January 1955 by north to south areas. found mainly in the neritic areas adja- chromic acid solution per 50 ml. of the cent to major river drainages. The sample. So far this fixative has been slope of the West Coast Florida con- the most effective in preserving G. breve tinental shelf is very gentle, so that a and other phytoplankton. depth of 60 feet lies 25-45 miles off- "Fixed water samples were stored shore; consequently, the neritic zone is under refrigeration at 36° F. These broad. samples were later filtered under re- "During periods of relatively light duced pressure, using HA gridded nnilli- concentrations, tremendously dense pore filters. After serial washes and sampling is necessary to validate the the application of a mordanting solution, presence of the organism. For ex- the organisnns were stained on the annple, on two different occasions, sam- millipore filter pad. The 3 most prom-

ples taken every hour in the same ising stains have been 1 percent Fast water mass for periods of twenty-four Green FCF, 1/2 percent Gentian Violet, hours showed the organism present in and 1/2 percent Crystal Violet. These but 3 out of 50 samples in one instance, filters are first cleared in cedar oil and in but 2 out of 48 in the other. Even or Cargille oil, then cut in half, and during periods of heavy concentrations mounted with balsam or Permount on a associated with fish kills, when sam- glass slide. A permanent slide index ples were in an area of dying fishes, of the phytoplankton taken since Janu-

the counts varied from to over 1 ary 1959 at selected stations is being million per quart, further indicating maintained. There is some tendency the extreme patchiness of the orga- for G. breve cells to round up during nism's distribution in sea water." filtration, but no distortion or cytolysis Two series of hourly samples taken of armored dinoflagellates or diatoms over 24- hour periods tend to show that was noted. the organisnn appears to concentrate at "This technique can be used to the surface during daylight and to move supplement living counts of G. breve downward at night. and associated phytoplankton and nnay be valuable during extended offshore FINUCANE, JOHN H. sampling trips when large changes in 1959a. Field ecology relating to red tide. the number of organisms could occur In Galveston Biological Laboratory fish- before examination of the samples." ery research for the year ending June 30, 1959, p. 76-79. U.S. Fish Wildl. FINUCANE, JOHN H. Serv., Circ. 62. 1960. Field ecology relating to red tide. In Galveston Biological Laboratory fish- "Since October, 1958, the dominant ery research for the year ending June phytoplankton associated with G. breve 30, 1960, p. 52-54. U.S. Fish Wildl. in the water samples were recorded Serv., Circ. 92. qualitatively in the inshore and off- shore waters of Tampa Bay. Quantita- "The 1959 red tide outbreak started tive counts (commenced in January 1959) during the last part of September and of dinoflagellates, diatoms and algae, continued through November. The initial made from preserved organisms stained pattern of distribution was similar to the on millipore filter paper, will be used 1957 outbreak with blooms of G. breve to supplement these qualitative data," first occurring outside the islands. The [p. 79.] first was reported September 29, offshore from St. Petersburg Beach. FINUCANE, JOHN H. The greatest observed numbers of 1959b. Fixing and staining of dinoflagel- G. breve (480,000-1,330,000 per liter) lates. &i Galveston Biological Labora- were found in the neritic waters 3-10 tory fishery researchfor the year ending miles west of Egmont Key, Septenaber June 30, 1959, p. 108. U.S. Fish Wildl. 30. By October there was a rapid de- Serv., Circ. 62. cline in both numbers and incidence of

G. breve , 10-40 miles west of Egmont "Due to the extreme fragility of Key. This was particularly noticeable

living G. breve , we are experinnenting at stations 10 miles offshore where the on techniques for the preservation of numbers dropped from 1,120,000 per plankton samples in the field which will liter in September to less than 100 per permit more precise quantitative and liter during October. The coastal waters qualitative analyses of samples. north of Egmont Key to Clearwater had "Field samples were fixed imme- a high count of only 33,000 per liter diately after collection with 0.5 ml. of as compared to 540,000 per liter at the 2 percent osmic acid in 1 percent stations south of Egmont Key to Venice

33 the same month. The last recorded fish above 30° C. n-iay inhibit blooms, kill during this outbreak occurred Octo- [p. 53.] ber 22, 1959, 3 miles west of Egmont Key. [p. 52.] "The incidence of G^. breve decreased in Tampa Bay and the nearshore waters "Blooms of Prorocentrum micans and during November, but blooms of this Ceratium furca occurred during Novem- organism to 400,000 per liter still oc- ber and December 1959 in Tampa Bay. . . curred 10 miles west of Egnnont Key. blooms of Gymnodinium splendens were Following the advent of weather fronts present March 29, I960, in upper and accompanied by lower water tempera- middle Tampa Bay. . . . tures and turbulent sea conditions, " Skuiaella erythraeum continued to be G. breve incidence and numbers de- the dominant blue-green alga in the clined during December 1959 and Jan- surface waters from Tampa Bay to 40 uary 1960. nniles west of Egmont Key during the "From February to April 1960 summer and fall of 1959." [p. 54.] G. breve increased again in numbers and distribution. In March, 35 miles FINUCANE, JOHN H. west of Egmont Key, a bloom of 6,320,000 196l. Ecology of red tide, hi Galveston per liter was observed. Scattered dead Biological Laboratory fishery research fish extended 15-35 miles offshore. No for the year ending June 30, 1961, further fish kills were reported in April p. 61-64. U.S. Fish Wildl. Serv., Circ. and May. The highest population en- 129. countered at that time was 21,000 per liter 10 miles west of Egmont Key. "A minor red tide developed in the neritic waters off Tampa Bay during

July and August 1 960. Blooms of G. breve as dense as 420,000 per liter were found "The neritic nature of G. breve was off Egmont Key, July 27, 1960. Fish indicated by its presence to depths of kills, principally confined to that area, 123 feet in numbers ranging from 7, 000- were of a very linnited nature. During 16,1 00 per liter, 40 miles west of Egmont August the greatest numbers of this Key, November 1959 and March 1960. organism (up to 150,000 per liter) were "Organisms were present inthe near- present fronrx 5 to 20 miles off Egmont shore and offshore waters throughout the Key. The number of samples containing year during non-bloom periods.... G. G. breve increased from a low of 3 per- breve was completely absent from the cent in June to 64 percent in August middle of Tampa Bay except during the during this outbreak. No further fish blooms of September-October 1959 and kills were recorded after the first week

February-March 1960. This suggests in August. . . ." [p. 61.] that G. breve is primarily a neritic organism found in estuarine waters only FINUCANE, JOHN H. when special conditions exist. 1964. Distribution and seasonal occurrence "Organisms were generally present of G ynnno d inium breve on the west coast in lower than normal salinities, ranging of Florida, 1954-57. U.S. Fish Wildl. from approximately 30-35^00 during Serv., Spec. Sci. Rep. Fish. 487, blooms. Waters having values less than iv + 14 p. 24^0 did not contain blooms of G. breve in Tampa Bay. While salinity may have "The tern-i 'bloom' represents any served as a barrier for G. breve in concentration of G. breve exceeding estuarine waters, it does not normally the normal population level of approxi- seem to be a limiting factor in neritic mately 1,000/1. G. breve counts of waters. more than 250,000/1. were considered "Water temperatures above 22° C. lethal during red-tide fish kills." [p. 1.] seem to be favorable for blooms of The coast from Tarpon Springs to G. breve with the optimum around Cape Sable is divided into 3 sectors. 26°-28° C. However, during February Tables and figures give the relative and March 1960, dense populations of abundance of G. breve in the samples G. breve developed at temperatures by sector by 2-month periods from between 15°-li C. Since G. breve January 1954 through December 1957. has been observed in waters as cold as In sector A (Anclote Keys to Venice) 9° C., low water temperatures nor- the organisms were just approaching mally may not be an absolute limiting bloom abundance during 1954, scarce factor for the existence of this orga- through 1955 and 1956, and in full nism. Temperatures below 14° C. and bloom during the last 4 nnonths of 1957.

34 .

In sector B (Venice almost to Naples) mortality of marine animals during the the organisms approached bloom during April 1963 red tide outbreak in Tampa the first half of 1954 and bloomed from Bay, Florida. Fla. Bd. Conserv., Mar. July 1954 through February 1955, then Lab., p. 97-107. remained scarce until they bloomed again during the last 4 months of 1957. Lethal concentrations of G. brev£ oc- In sector C the organisms showed a curred for about 2 weeks, April 3-16, sudden increase to just below bloom 1963, in an area from the mouth to about level from March through June 1954, 25 n-iiles inside Tampa Bay. No blooms and then commenced blooming and con- were detected in upper Hillsborough through tinued February 1955. Bloom Bay or Old Tampa Bay. Fifty-six species occurred again during the last 4 months of fish were killed, also blue crabs, of 1957. horseshoe crabs, and lancelets, but In than samples, breve more 9,000 G. dead shrimp were not seen. The authors not found in salinities below 21 or was studied seine and trawl hauls made in over 39 p.p.t., or at temperatures be- the area in 1962 and 1963 and concluded low 10° or above 34° C. that the effects of red tide on fish In simultaneous surface and bottom populations appear to be rather limited. samples G. breve was higher in the surface samples. The great majority of the samples were collected, how- FISH, CHARLES J., and MARY CURTIS COBB, ever, between 8 a.m. and 5 p.m. 1954. Noxious marine animals of the central

". . . Since its incidence during non- and western Pacific Ocean. Fish Wildl, bloom periods is primarily confined to Serv., Res. Rep. 36:iii + 45 p. offshore waters, the organism prob- ably is more neritic than estuarine. Includes a few references onredtide. G. breve occurred in approximately equal frequency in both estuarine and FISHER, A, neritic waters only during the red- 1956. The effect of copper- sulphate on some tide outbreaks in 1954 and 1957. The microorganisms in fish ponds. offshore distribution usually extended Bamidgeh, Bull. Fish Cult. Israel 6-10 miles, although in March 1960 8(2):21-27. a bloom of 6,320,000/1. was detected miles offshore (Hutton, 1960). During 35 Daphnia and Cyclops were not harmed the red-tide outbreak of 1954, Lackey by 1 mg,/l. of copper sulphate; at and Hynes reported G. breve (1955) 10 mg./l, Daphnia lived 35 hours. in samples collected 140 miles south- Cyclops was less affected than Daphnia . west of Fort Myers, Fla. . . ." [p. 8.] FLORIDA STATE BOARD OF CONSERVATION. FINUCANE, JOHN H. and ALEXANDER Conserv., DRAGOVICH. 1959. Red tide. Fla. State Bd. 13th Bien. Rep. 1 957- 1 958, p. 1 1 1959. Counts of red tide organisms, Gym-

nodinium breve , and associated ocean- A general account of efforts to con- ographic data from Florida west coast, trol the 1957 red-tide outbreak. 1954-57. U.S. Fish Wildl. Serv., Spec, Sci. Rep. Fish. 289, iv + 220 p. FORTI, A. 1933. 11 fenomeno de "lago di sangue" nello This report contains the original stagno di Pergusa in Sicilia alia meta data on counts of the red-tide orga- di Settembre 1932, Nuovo Giorn. Bot. nism, G. breve , and concomitant physi- Ital, 40(l):76-78, cal and chemical hydrographic data collected from February 1954 through June 1957, between Clearwater and FRITSCH, F, E. Marathon Key on the west coast of 1935. The structure and reproduction of the Florida. Hydrographic data include algae. Vol. I. Univ, Press, Cannbridge, temperature, salinity, pH, copper, in- 791 p. organic and total phosphorus, nitrate- nitrite, and carbohydrate and protein Contains 176 references on Dino- equivalents. phyceae.

H., GORDON R. RINCKE\ GALTSOFF, PAUL S. FINUCANE, JOHN , and CARL H. SALOMAN, 1948. Red tide. Progress report on the

1 964. A collection of data in reference to red investigations of the cause of the mor- tide outbreaks during 1963. 5. Mass tality of fish along the west coast of

35 Florida conducted by the U,S, Fish thousands were washed onto the

and Wildlife Service and cooperating beaches . , . [p. 13-14.] organizations. Fish Wildl. Serv,, Spec. Sci. Rep. 46, i + 44 p.

During the 1946-47 outbreak: All kinds of animals perished in the red "... the water had an oily appear- water, including a small number of ance. When dipped up and allowed to turtles and porpoises. Windrows of stand for 5 to 1 minutes, it became dead fishes piled on beaches com- thick, sometimes almost of a con- prised a great variety of connmon com- sistency of Karo syrup, and slimy to nnercial and noncommercial varieties. the touch. . . . [p. 15.] Likewise, the pelagic and bottom in- vertebrates succumbeS to the unknown poison. Large numbers of shrimp were seen dead, as well as common blue crabs, fiddler and mud crabs, barnacles, ". . . Counts made by Woodcock and oysters and coquinas. Observations Anderson in the field show that the made by the author in March 1947, around number of Gymnodinium in the surface the Fort Myers area, disclosed that layer of red water varied at this time about 80 percent of the oysters, Ostrea from 13,000,000 to 56,000,000 per liter.

virginica , grown on piles, were dead. Samples collected by plankton net con-

"No n-iortality was observed among tained also large numbers of Evadne . the hard shell clams, Venus mercenaria , The intestines of this Cladoceran were and no reports were received of the stained deep red by ingested Gym- destruction of ducks, gulls, and other nodinia. . . ." [p. 19.] birds inhabiting the inshore waters. Later during the summer infornnation was received fronn residents of Largo, Florida, that 'thousands of sea gulls and pelicans died from eating the fish ", . . Dinoflagellates, like other poisoned by the red tide.' The correct- , are very sensitive to copper ness of this observation has not been sulphate and hypochlorite. Kofoid and verified by the author." [p. 10-11.] Swezey (1921) state that copper sulphate

In June 1947, W. W. Anderson re- in a concentration of 1 part per million ported the species killed in the Fort killed all Ceratium hirundinella . This Myers area to be 50 percent catfish; method was used by the Japanese biolo- also included were pinfish, porgies, gists in their attempts to control the white trout and spotted trout, cowfish, red water in the Gokasho Bay and in spiny box fish, moonfish, spot, mullet, the Gulf of Konsa. Miyajima (1934) states eel, sand-bream, whiting, thread that all dinoflagellates are instantly herring, hogchoker, tongue fish, yellow- killed by a copper sulphate solution tail, tripletail, redfish, and drum. Also in the concentration of 2 parts per very noticeable were carcasses of million. The concentration of 1 part horseshoe crabs. per million kills thenn within a few

". . . Sponge divers working last winter nninutes. In practice the copper sulphate off Marco, Florida, reported that the was applied by attaching bags filled with bottom was littered with dead mackerel, this salt to the sides of motor launches although rarely were these fish found which were run back and forth in the on the beaches, [p. 13.] bay. After the treatment large numbers "Although few dead crabs were ob- of destroyed dinoflagellates were found served by Anderson, this may be due floating in the water, [p. 34.] to the fact that these animals tend to "To prevent the growth of bacteria sink upon death and would, therefore, which may develop after the destruc- be less noticeable. On the other hand, tion of dinoflagellates, the Japanese he frequently observed small species of biologists suggest the use of 1 percent crabs swimming in the infected waters solution of calcium hypochlorite or the in apparently good condition. On several addition of liquid chlorine. Both solu- occasions crabs were observed feeding tions can be used simultaneously and on dead fish at the surface. He thinks their effective concentrations, accord- that it is entirely possible that these ing to Kominarui (quoted from Miyajima) crustaceans were less affected by the should be adjusted to the salinity and red water than were the fish. Horse- temperature of the water. He states that shoe crabs (Limulus polyphennus ), how- at 13.6° C. ordinary bleaching powder ever, suffered a heavy mortality and containing from 34 to 35 percent of free

36 chlorine is effective in killing dino- Gonyaulax killed fish at concentra- flagellates at the concentration of tions of 1,400 per liter. The fish lost 1:500,000. At 10° C. the concentration equilibrium, then slowed their opercular should be increased to 1:400,000. In movements, combination with copper sulphate the concentration of hypochlorite can be GILSON, H, GARY, Murray reduced to 1 part per million. It is 1937. The nitrogen cycle, John interesting to note that after the red Exped,, 1933-34, 2{2):21-81, [Cited water was destroyed by chemical treat- from Ryther, 1955.] ment the Japanese biologists noticed the appearance of another dinoflagellate GLAZIER, W, C. W, of the genus Polykrikos which apparently 1882, On the destruction of fish by polluted was harmless to pearl oysters. waters in the Gulf of Mexico, Proc. Nat, 4:126-127, ". . . it appears promising to try the U,S, Mus, spraying of red water fron-i aeroplanes or from boats with a solution of copper Fish in live wells of fishing smacks sulphate or dusting it with powdered have often died when passing through calcium hypochlorite. Other chemicals, poisoned water, notably about 1865 and harmless to fish and shellfish, may be in 1878, when large numbers were tried. The use of powdered calcium thrown on the shore at Key West, oxide (unslackened lime) suggested it- He reported (writing on November 25, self, for its addition to sea water will 1880) that the waters of Tampa, Sara- raise the pH to a level which is beyond sota, and Charlotte Harbor were covered the tolerance of the dinoflagellate and, with thousands of dead fish, and several at the same time, it is unlikely that smacks lost their cargoes in the last the increased concentration of Ca salts 2 weeks, in the water will adversely affect fish or shellfish, for the excess of Ca in GLENNAN, A. H, sea water will be rapidly precipitated, 1887, 4, --Fish killed by poisonous water. "Advantage may be taken of the fact U,S, Fish Comm,, Bull, 6:10-11, that the greatest concentrations of dino- flagellate s appear in patches which Writing on October 28, 1885, Glennan are, probably, the centers of their stated that large shoals of dead fish Light more rapid propagation. . . ." [p. 34-35.] were seen between Egmont Key and Charlotte Harbor, He reported that a fishing schooner a few weeks earlier GALTSOFF, PAUL S. had lost two loads of live fish in sailing 1949. The mystery of the red tide. Sci. through strips of this poisoned water, Mon. 68 (2):108-117. GOODELL, H, G,, and D, S. GORSLINE. Semipopular account of information 1961, Data report - the hydrography of given in his 1948 report. Apalachicola and Florida Bays, Florida, Fla, State Univ., Sedimentological Res. GATES, JEAN. Lab,, Contr, 1:1-316. 1958. Toxicity of Gonyaulax nnonilata to fish. In Annual report of the Gulf Fish- GRAHAM, HERBERT W, ery Investigations for the year ending 1942, Studies in the morphology, taxonomy, June 30, 1958, U.S. Fish Wildl. Serv., and ecology of the Peridiniales, Car- 90-93. p. negie Inst., Wash,, Publ. 542, 129 p.

This material was published by Gates GRAHAM, HERBERT W. and Wilson (I960). 1954, Dinoflagellate s of the Gulf of Mexico. In Paul S, Galtsoff (coordinator). Gulf GATES, JEAN A., and WILLIAM B, WILSON. of Mexico: its origin, waters, and ma- 1960. The toxicity of Gonyaulax monilata rine life, p, 223-226. Fish Wildl, Serv,,

Howell to Mugil cephalus . Limnol. Fish, Bull, 55.

Oceanogr. 5(2): 1 71 - 174. "Toxic red water such as occurs Gonyaulax monilata caused yearly regularly in the pearl oyster beds in summer mortality in Offatts Bayou from Japan (Mitsukuri 1904) could be dis- 1936-1941 (Gunter, 1942), Connell and astrous to the vast oyster industry in Cross (1950) reported mortality "almost the Gulf, but apparently the Gulf oysters every year" for 15-20 years prior to have been spared any such visitation 1949. so far.

37 "Reports of red water on Campeche 2.35 ug.at./l. of total phosphorus and Banks, off Yucatan, are made occa- only 0.05 of inorganic phosphorus. These sionally by fishernnen in that area, but analyses indicated that the alga can to date it has not been possible to grow well in water containing scarcely ." and ascertain the causative agent. . . measurable inorganic phosphorus [p. 225.] accumulate a highquantity of phosphorus and that high values of phosphorus in GRAHAM, HERBERT W., JOHN M, AMISON, unfiltered samples of water containing and KENNETH T. MARVIN. red tide may have little bearing on the 1954. Phosphorus content of waters along need for any specific amount for a bloom. the west coast of Florida. Fish Wildl. The authors observed Skujaella float- Serv., Spec. Sci. Rep. Fish. 122, ing in bands several hundred yards V + 43 p. wide and miles long. They speculated that if conditions at any time suddenly Total, inorganic, and organic phos- became vinfavorable for Skujaella a phorus was determined for waters of great mass of organic matter would the Peace and Caloosahatchee Rivers, beconne available to decompose and Charlotte Harbor, and 10 stations in release large quantities of nutrients the Gulf of Mexico off the west coast that nnight cause blooms of other orga- of Florida extending to 120 miles off- nisms.

shore. The period covered was May 1 949 Their data show that the phosphorus- to January 1951, inclusive. Water sam- rich waters of the Peace River do not ples were also analyzed for salinity, normally increase the phosphorus con- oxygen, and pH. This report contains tent of local Gulf water to any measurable only the phosphorus data; the rennainder degree. The phosphorus in the surface were published by Marvin (1955). waters of the Gulf gradually decreases The authors stated, "No high degree out to a distance of about 85 miles. of accuracy is claimed for the analyses Phosphorus found in larger quantities of the river water, especially at Station at depths below 50 m. was largely 2 (Peace River). Concentrations there inorganic. Limited upwelling of deep were frequently so high that dilutions water at certain times had no apparent with salt solutions were necessary. effect on the phosphorus content of Furthermore, yellowish or greenish water in the euphotic zone. tints often developed in the samples There was no evidence that bottom so that addition of dyestuffs to the sediments contributed appreciable quan- standards was necessary in order to tities of phosphorus to Gulf waters. effect a match in color. "In an effort to clarify the water, GRAHAM, HERBERT W., and NATALIA Sonne river samples were centrifuged. BRONIKOVSKY. It was found that not only the total, 1944. The genus Ceratium in the Pacific i>ut also the inorganic phosphorus was and North Atlantic Oceans. Carnegie less in these samples, indicating that Inst., Wash., Publ. 565, 209 p. some of the inorganic phosphorus oc- higher curs in particulate form. . . ." [p. 2.] Mentions the tendency toward a The authors pointed out the possible nunnber of species in areas lower in

role of Skujaella ( Trichodesmium ) in ,

concentrating phosphorus. ". . . This filamentous blue-green alga is always GRAN, H. H. present in the plankton there and fre- 1929. Investigation of the production of quently occurs in bloom proportions. plankton outside the Romsdalsfjord It grows in the water and on the sur- 1926-1927, 112 p. Cons. Perma. Int.

face. . . . Its propensity for floating on Explor. Mer, Rapp. Proc.-Verb. R^un. the surface is the feature which is of 56. [Cited from Ryther, 1955.]

importance to the present problem. . . . "Since it accumulates at the surface GULF COAST SHELLFISH SANITATION RE- it is driven by the wind. Sometimes it SEARCH CENTER. piles up in such abundance as to create 1964. Adverse chemicals and in the ." a nuisance at bathing beaches. . . marine environment and shellfish. U.S. [p. 40.] Public Health Service, Shellfish Sanit. In four samples of water containing Res. Planning Conf. Feb. 25-26, 1964.

blooms of Skujaella , the inorganic phos- Dauphin Island, Ala,, 9 p.

phorus ranged from 0.05 to .20ug.at./l ., and the total phosphorus from 1.75 to In December 1962 several illnesses 10.20 ug.at./l. A filtered portion of the reported to the Sarasota County Board sample with 10.20 ug.at./l. showedonly of Health were thought to have been

38 )

caused by eating shellfish. The Florida "In the summer of 1935 a very heavy State Board of Health made bioassays mortality of marine fishes and other for toxicity on shellfish san-iples for animals occurred in the Gulf of Mexico several southwest coastal counties. and extended for a distance of 250 Since May 1963 the Florida State Board miles from the Rio Grande northward. of Conservation has been identifying A few fishes were killed in the lower and counting phytoplankton in water bays but practically all of the mortality samples collected from the sanae areas was confined to the Gulf, The amount as the shellfish samples. of destruction was undetermined, but Since May 1963 the U.S. Public Health it was certainly enormous and ran into Service Laboratory (Dauphin Island) has many million pounds. Since that time been collecting shellfish samples from there was one other occurrence of the St. Petersburg-Sarasota area for this phenomenon in the fall of 1948. bioassay of toxicity. Since September It occurred along the southern tip of ." 1963 regular shellfish samples have the coast and was more localized. . . been taken in Lemon Bay, [p. 65,] "From the beginning many workers have considered that shellfish toxicity GUNTER, GORDON, F. G. WALTON SMITH, was casually [sic] related to the occur- and ROBERT H, WILLIAMS. rence of Red Tide. The work so far 1947, Mass mortality of marine animals has shown no results that are contra- on the lower west coast of Florida, dictory to this relation. In fact, many November 1946- January 1947. Science of the results suggest that the relation 105(2723):256.257. is true. However, rigorous proof of the Red Tide organism as the cause Dead fish were reported from Dry of shellfish toxicity remains to be Tortugas to Boca Grande (130 miles). demonstrated." [p, 7.] At Fort Myers dead fish exceeded 170 per foot of shoreline on January 19. GUNTER, GORDON. Total number of dead fish was esti- 1947. Catastrophism in the sea and its mated at 50 million. Also killed were paleontological significance, with oysters, clams, crabs, shrimp, barna- special reference to the Gulf of Mexico. cles, and coquinas. The clam industry Amer. J, Sci, 245(11 ):669-676, at Marco, 50 miles south of Fort Myers was not involved. Despite an isolated fish a vessel ". , , In connection with a heavy fish report of dead from mortality on the Texas Coast, Lund passing Dry Tortugas, mortality did (1936) called attention to the possible not appear to reach the Keys, A patch similarity of the case of Jordan's (1921 of vivid yellow water south of Useppa description of 'A Miocene Catas- Island on January 28, consisted alnnost trophe,' ... [p. 669-670,] exclusively of Gymnodinium sp. with a mixture of numerous larval inverte- brates. The water was viscid and slimy, "Mass mortalities of marine animals having the consistency of diluted syrup. on the west Florida Coast have been The oxygen was low, 33 percent satu- reported by Pierce (1883), Taylor (1917) rated. Irritating gas from breaking surf and Gunter, Williams, Davis and Smith occurred January 22-26 on Captiva (1947), According to Taylor (1917) this Island. phenomenon has been reported since 1844. It came again in 1946 and 1947. GUNTER, GORDON, ROBERT H. WILLIAMS, It has appeared on the average of CHARLES C, DAVIS, and F. G. WALTON

10.4 years since 1 844 with skips ranging SMITH, from one to thirty years. Occurrences 1948, Catastrophic mass mortality of ma- in consecutive years such as 1883-84 rine animals and coincident phyto- and 1946-47, are continuations of the plankton bloom on the west coast of Florida, November 1946 to August 1947, same outbreak. . . ." [p. 673.] Ecol, Monogr. 1 8(3):309-334, GUNTER, GORDON, 1949. The "red tide" and the Florida fish- The mortality was first noticed No- eries. Proc. Gulf Carib. Fish. Inst., vember 20, 1946, 14 miles offshore Inaugural Sess.:31-32. from Naples, It spread as far north as Boca Grande Pass. According to all GUNTER, GORDON. reports, no mass mortality occurred 1952. The import of catastrophic mortalities north of . Mortality for marine fisheries along the Texas occurred southward definitely as far as coast. J. Wildl. Manage. l6(l):63-69. Cape Romano. On January 19 the captain

39 .

of a small freighter reported dead east coast of Cape Peninsula was caused fish from Dry Tortugas to Fort Myers, by a ciliate he thought belonged to the

After the end of January 1947, no genus Mesodinium . further reports were received of dead fish until April 2, when they were HART, T. JOHN. reported from Cape Florida to Mara- 1942. Phytoplankton periodicity in Antarctic thon and west almost to Key West. surface waters. Discovery Rep. Later fish mortality was reported south 21:261-356. of the Keys in some places but appar- ently it had disappeared by the end of HASLE, GRETHE RYTTER. April. Total area covered by the out- 1950. Phototactic vertical migration innna- break was from Sarasota to the Florida rine dinoflagellates. Oikos 2:162-175. Keys, and duration was from November 1946 to the latter half of August 1947. HASLE, GRETHE RYTTER, Mortality occurred in turtles, bottle- 1954. More on phototactic diurnal migration

nose ( Tursiops truncatus ), Bala- in marine dinoflagellates. Nytt Mag.

nus , oysters, coquinas Donax Bot. 2:139-147. [Cited from Pomeroy ( ), Penaeus ,

crabs ( Callinectes sapidus most com- et al., 1956.] mon), and about 25 species of fishes. The estimate of 50 million fish killed HAXO, F. T., and BEATRICE M. SWEENEY. (Gunter, Smith, and Williams, 1947) 1955. Bioluminescence in Gonyaulax polye- was revised to 500 million. The num- dra. In Frank H. Johnson (editor). The ber of dead fish averaged about one luminescence of biological systenns, per square yard up to 4 or 5 miles off- p. 415-420. Amer. Ass. Advance. Sci., shore and about one per acre 10 miles Washington, D.C. offshore. The average maximum air tempera- Gonyaulax polyedra may need other ture at Fort Myers for the 4 months vitamins in addition to Bj2> thiamine, of October 1946 through January 1947 and biotin. was abnormally warm- -6.0° F. above the 55- year average. The hurricane of HAYES, HELEN LANDAU, and THOMAS S. October 7, 1946, might also be impli- AUSTIN. cated. 1951. The distribution of discolored sea water. Tex. J. Sci. 3(4):530-541 GUNTHER, E. R. 1936. A report on oceanographical investi- Contains 225 references pertaining gations in the Peru coastal current. to the occurrence and causes of the DiscoveryRep. 13:107-276. [Citedfrom overblooming of plankton, with special Pomeroy et al., 1946.] emphasis on dinoflagellates.

Upwelling of water into photic zone HELA, ILMO. is given as a source of nutrients. 1955. Ecological observations on a locally limited red tide bloom. Bull. Mar. Sci. HALE, FRANK E. Gulf Carib. 5(4):269-291. 1950. The use of copper sulphate in control of microscopic organisms. Phelps The study was made to determine the Dodge Refining Corp., New York, 43 p. effect of passes in generating red-tide outbreaks. On p. 270 it is stated, "Thus HARRISON, EDWARD F. it was possible to perform observations 1957. A study of the so-called red tide in under the actual conditions of an initial the Gulf of Mexico. Privately printed, Red Tide outbreak." The sampling was 6 p. apparently done only from November 30 to December 2, 1954, however, Suggests controlling red tide by dyna- and the outbreak was under way some- miting the nnud lunaps off the mouth what earlier. At their station B (Station of the Mississippi River and then vio- B29 in Finucane and Dragovich, 1959) lently agitating the water with water counts of G. breve were 30,000 per jets to liberate trapped gases. liter on November 24, and at their station X5, where they counted 60,000, HART, T. JOHN. the count on November 24 was 140,000. 1934. Red "water-bloom" in South African The author attempted to explain the seas. Nature 134(3386):459-460. patchiness of G. breve concentrations by stating that the ebb tide flows westerly In nnid-July 1934 a blood-red dis- (actually southwesterly) down Boca coloration along several nniles of the Grande Channel, and suggested that the

40 flooding waters probably come from end of which seems to be at the latitude the region south of the Pass and cut of Tampa Bay. off the ebbing waters. He stated,

". . . Thus this mechanism indicates HIRASAKA, KYOSUKE. how the coastal water patches may be 1922. On a case of discolored sea-water.

formed in the narrow pass by the flood Annot. Zool, Jap. 1 0( 1 5): 1 6l - 1 64. current completely cutting off each [Cited from Martin and Nelson, 1929.] ebbing water movement." [p. 275.] The author attempted to show these According to Martin and Nelson (1929), patches in figure 12 by imaging four Hirasaka stated red water caused by successive patches of water in a row Gymnodinium sanguineum did no dam- of eight G^. breve observations extend- age, but he cited an earlier outbreak ing altogether about 6 miles down Boca that injured some fish, which he re- Grande Channel. In the face of the 2- ported from Okamura; and also from knot currents he showed this conclusion Kofoid and Swezy (1921). to be unreasonable, as four patches HOLLAND E, ANDRE, and MONIQUE would take four tidal cycles to form. ENJUMET. Additionally, the slightly lower counts 1957. Sur une invasion des eaux du port at his interpatch stations probably are d'Alger par Chattonella subsalsa not statistically significant. (= Hornellia marina sub.) Biecheler. We do not agree with the optimum Remarques sur la toxicite de cette salinity for G. breve suggested the by Chloromonadine. Bull. Sta. Aquic. points plotted in figure 15. An attempt Peche Castiglione^n.s,,8(1965):273-280. was made to prove vertical migration

in G. breve ; however, the data presented HORNELL, JAMES. do not afford adequate proof, 1917. A new protozoan cause of widespread mortality among marine fishes. Madras HELA, ILMO, DONALD deSYLVA, and Fish. Dep., Bull. ll(2):53-66. CLARENCE A, CARPENTER. 1955. Drift currents in the red tide area "Widespread fish mortality is a well of the easternmost region of the Gulf known phenomenon on the Malabar and of Mexico. Univ. Miami, Mar. Lab., South Kanara coasts; its recurrence Rep. to Fla, State Bd. Ccnserv. 55- yearly along certain stretches of the 11. 31 p. coast line is regular, though its intensity

During 1954 two driftcard operations varies within wide limits. . . . were carried out off the west coast of Florida. The ultimate goal was to solve

the problem of the peculiar nearshore ". . . all Malabar fishermen whom I movements of the floating dead fish. have questioned agree in saying that "It was assumed that under the com- every year after the passing of the rainy bined influence of the (1) more or less season and the subsidence of the south- permanent ocean currents and of the west monsoon, if there be a continuance

(2) highly variable drift currents , pro- of fine weather for a week or ten days, duced locally by the winds, the paths with plenty of sunshine, and a weak of the driftcards would be roughly the coastal current, the water inshore be- same as the paths of the dead fish comes turbid and discoloured, brownish patches. In this report it is shown or reddish in tint; that this water has that the estimated path of the drift- such poisonous effects upon fish that cards, and therefore the drift of dead large numbers become affected and

fish patches, can for practical pur- eventually die. . . ." [p. 53.] poses be determined by means of the

wind observations alone. . . ." [p. 1.] HORNELL, J., and M. R. NAYUDU. On July 13, 1954, a total of 1,100 1923. A contribution to the life history of cards were dropped, 10 at each mile the Indian sardine. Madras Fish. Dep,, along four transects, 240° true, up to Bull. 17:129-97. [Citedby g.yther, 195 5. 30 miles offshore, between San Carlos Non vidi.] Bay and Longboat Inlet. Another 1,000 cards were dropped in 18 passes, from Ryther stated the report described Wiggins Pass north to Palma Sola, On red water caused by peridineans occur- November 17, 1954, 2,200 cards were ring annually along the Malabar Coast, dropped offshore by plane, in the same following the cessation of the southwest general area. monsoon, diatom blooms, and heavy The authors stated the returns showed rainfall. Blooms occur during hot, calm the existence of a permanent counter- weather in bands along the shore or in clockwise current eddy, the northern patches which nnove with the tide.

41 HOWELL, JOHN F. They showed a photomicrograph of

1953. Gonyaulax monilata , sp. nov., the Zooxanthellae, symbiotic algae that causative dinoflagellate of a red tide live in jellyfish and coral. When these on the east coast of Florida in August- simple round cells were grown in proper September, 1951. Trans. Amer. Micro- media, they developed into the dinoflag- scop. Soc. 72{2):153-156. ellate Gymnodinium adriaticum, mobile and with flagella. Contains a description of Gonyaulax

monilata . Author said that it killed HUTNER, S. H., L. PROVASOLI, ALBERT fish, but no great quantity, in Indian SCHATZ, and C. P. HASKINS. River, Fla., during August 1951. 1950. Some approaches to the study of the role of metals in the metabolisn-i of HUTCHINSON, G. EVELYN. microorganisms. Proc. Anner. Phil, 1944. Limnological studies in Connecticut. Soc. 94(2):152-170, VII. A critical examination of the sup- posed relationship between phytoplank- HUTTON, ROBERT F. ton periodicity and chemical changes 1956. An annotated bibliography of red tides in lake waters. Ecology 25(l):3-26. occurring in the marine waters of [Cited from Ryther, 1955.] Florida. Quart. J. Fla. Acad. Sci. 19(2-3):124-146. HUTNER, S. H., and JOHN J. A. McLAUGHLIN. 1958. Poisonous tides. Sci. Amer. 199(2): "At Ballast Point in Hillsborough 92-96, 98. Bay south of Tampa, Florida, a minor Red Tide occurred periodically during A general discussion of red tides. 1955. Fishes did not seem to be harnned

Dinoflagellate toxins appear to block as minnows ( Gambusia ) and mullet

nerve impulses by preventing the pro- ( Mugil ) were seen swimming through the duction of acetylcholine, the substance streaked areas from time to time. which acts on the "end plate" of a muscle However, shrimp in bait-wells located fiber to make the fiber contract. This on the fishing pier at Ballast Point died is sinnilar to the action of botulinum, when this water was pumped into the the toxin of botulinus. No antidote to wells although good water circulation either poison has been found. was maintained. A dinoflagellate, "The prospect for restraining blooms Ceratium furca (Ehrenberg), was the by removing essential nutrients from predominant organism in the water the water seems poor. Vitamin B-12, exan-iined and counts as high as for example, could conceivably be cut 17,600,000 cells per liter were ob- down by dumping carboys of B-12- served." [p. 125.] destroying bacteria in the water, but The author stated that, "Lackey (1 955) truly enormous quantities of bacteria reported a bloom of the diaton-i Coscino- would be required to make any im- discus in the area around the nnouth pression on several hundred square of the Myakka River in 1953." [p. 125.]

miles of ocean. . . A nnore promising The indicated reference, however, is attack seems to be the encouragement not listed. of natural predators, the technique which The annotations are largely uncritical; has so often been successful with insect they serve as a guide to contents and pests. Dinoflagellate blooms are often often consist of the author's own ab- succeeded by blooms of creatures that stract. prey on them, most conspicuously the ciliate protozoa and, in warm waters, HUTTON, ROBERT F. the luminous, I960. Notes on the causes of discolored

Noctiluca. . , . [p. 95.] water along the southwestern coast of Because B-12 and the pseudo-B-12 Florida. Quart. J. Fla. Acad. Sci. 23(2): vitamins are large molecules built 163-164. around an atom of cobalt, they sug- gested there may be unproductive areas Discoloration can be caused not only of the ocean caused by lack of cobalt, by red tide but also by a blue-green - similar to areas on land known as alga, Skuja^lla ( Trichodesmium ) thie "cobalt-deserts" where livestock die bauti De Toni, which causes discolora- from lack of cobalt in the soil. tions frequently reported by boat cap- They stated that in fresh water the tains and airplane pilots as red tide. growth of Euglena is a sensitive indi- During 1960 discolorations had three cator of the amount of B-12 present, causes:

but no similar organism has been dis- 1 . Gymnodiniunn breve Davis. In Jan- covered for use in salt water. uary 1960, counts were as high as 7

42 million per liter between Cape Romano it continued to blow from the south ." and Englewood. On March 23, counts and west until the 11th of October. . . were over 6 million per liter 15 to 35 [p. 77.] miles west of Egmont Key. A report containing raw hydrographic 2. Gynnnodinium s pi end ens Lebour. data obtained in an area north of Key On March 27 several square miles were West from Cape Sable to Dry Tortugas discolored south of Gandy Bridge in {July-December, 1961). Tampa Bay with counts of more than 5 nnillion per liter (no apparent damage INGLE, ROBERT M. to fish). 1954. Irritant gases associated with red 3. Acartia tonsa Dana and Labidocera tide. Univ. Miami, Mar. Lab., Spec. aestiva Wheeler. These Crustacea Serv. Bull. 9, 4 p. caused a streak of reddish brown water 1 1/2 miles west of Big Sarasota Pass Gives blooming of Gymnodinium on March 29. No dead fish were seen. brevis as the cause of irritant gases on Counts of A. tonsa were over 350,000 beaches of Florida's west coast. The per liter, but L. aestiva was much less author said that gases are given off abundant. only if red-tide waters are heated or agitated. The theory is discounted that ICHIYE, T. the poisonous gases bring about growth

1962. Circulation and water mass distribu- of G. brevis . Irritant effects are only tion in the Gulf of Mexico. Fla. State temporary and "do not appear unless Univ., Oceanogr. Inst., Contr. 190, 76p, wind-driven waves with associated water vapor and droplets exist." [p. 1.] INGERSOLL, ERNEST. 1882. On the fish-mortality in the Gulf of INGLE, ROBERT M., and DONALD P. Mexico. Proc. U.S. Nat. Mus. 4:74-80. deSYLVA. 1955. The red tide. Fla. State Bd. Conserv., He attempted to discover facts con- Educ. Ser. 1, 30 p. [Revision of 1948 cerning fish kills that had occurred in edition.] 1880 and came up with the following information from talking with residents: For general information of the public. 1) As far back as 1844 [in 1844?] a widespread destruction occurred. INGLE, R. M., R. F. HUTTON, H.E.SHAFER, 2) This occurred again in 1854. JR., and R. GOSS. 3) This has occurred at intervals 1959. The airplane as an instrument in since, to a less degree. marine research. Part I. Dinoflagel- 4) In 1878 an "excessive fatality" late blooms. Fla. State Bd. Conserv., occurred, probably more destructive Spec. Sci. Rep. 3, 25 p. than the later visitation of 1880. He placed so much emphasis on the death The authors reported that semicircu- of sponges in 1878 that one wonders lar surface structures were common whether the 1878 incident was red tide features of the 1957 red-tide outbreak or a sponge disease. near the mouths of passes. "The muddy

5) Even the cooler half of 1879 was bay water . . . lay proximal to the pass. not exempt from some appearance of Dead fish were not abundant in the the plague. muddy less saline water. Typically, the 6) The 1880 red tide appeared in interface between the water masses was August immediately following "the filled with a line of dead fish. [p. 3.] terrible hurricane which is known as the 'August Gale.' " Fish died along the southern shore of Tampa Bay and "In every case observed by the au- Egmont Key south to Shark River, thors (R.M.I, and R.F.H.) thediscolored south to Bahia Honda Passage, and water, denoting a high concentration of beyond Key West. dinoflagellates, was found from the in- 7) At Egmont Key the first dead fish terface and extending seaward. occurred on October 17, 1880, and the "Because of their role in concentrat- red tide was in full swing by October ing nutrients and dinoflagellates, and 25. Fish continued to die for 6 weeks, due to their frequency of occurrence, at a decreasing daily rate. interfaces may well serve to rejuvenate

". . . there was immense quantities and sustain red tides once they begin. If of fresh water coming down the bay, this is true, control measures directed and the water here [Egmont Key] was to the specific areas of junction of dis- nearly fresh on the surface, while the similar water masses may serve a

water underneath was perfectly salt. . . . useful purpose." [p. 5.]

43 This report states that after October tion must have been very great, for 15, 1957, spraying of copper sulfate here, on a key containing but a few crystals was discontinued and control acres, and with a very limited extent measures consisted of suspending bags of beach, we have buried at least

of copper sulfate off bridges, etc. in 27 twenty cart-loads, , , ." [p, 244.] passes between Clearwater and Naples. The letters also alluded to fish dying in and to death of almost INGLE, ROBERT M., and JAMES E. SYKES. all the conchs around the Dry Tortugas. 1964. A collection of data in reference to red tide outbreaks during 1963. 1. 1963 JOUAN, H. ^ red tide and associated studies--a pre- 1875. Melanges zoologiques. Mortalite' sur liminary report. Introduction. Fla. Bd. les poissons a la c6te de Malabar. Conserv., Mar. Lab., p. 2-3. Mem. Soc. Sci. Nat., Cherbourg 19:233- 245. An introductory paper to several re- ports on an April 1963 outbreak of red JUNGS T, H. tide in Tampa Bay. 1937. Fischsterben im Kurischen Haff, Geol, Meere Binnengewasser 1:352-354. JEFFERSON, J. P. 1879. On the mortality of fishes in the Gulf KAISER, E, of Mexico in 1878. Proc. U.S. Nat. 1930, Das Fischsterben in der Walfisch- Mus. 1:363-364. bucht. Palaeobiologica 3:14-20,

In his letter (written in December KETCHUM, BOSTWICK H,, and JEAN KEEN, 1878) to Spencer F. Baird, hedescribed 1948. Unusual phosphorus concentrations a second occurrence (see Jefferson et in the Florida "red tide" sea water. al., 1879) of a large body of dark- J. Mar. Res. 7(1):17-21. colored water moving southward along the Florida coast, across Florida Bay, Several sea water samples were taken and striking the Tortugas about Novenn- in July and August 1947, preserved with ber 20, 1878 and extending up the reef chloroform, and shipped to Woods Hole as far as Key West, probably farther. for analysis. Their phosphorus values He mentioned that oysters in Tampa include the organic phosphorus com- Bay were killed by the water. He also bined in particulate matter and plankton, reported that in October the Caloosa- the organic phosphorus in solution, and hatchee River overflowed its banks inorganic phosphate-phosphorus. along its entire length, except at a bluff High total phosphorus values were in Fort Myers. obtained in the samples containing heavy blooms of red tide. The only way the JEFFERSON, J. P., JOSEPH Y. PORTER, authors could account for the high con- and THOMAS MOORE. centration was by assuming that the 1879. On the destruction of fish in the organisms were able to accumulate the vicinity of the Tortugas during the phosphorus from the entire water col- months of September and October, 1878. umn. Proc. U.S. Nat. Mus. 1:244-246. KIERSTEAD, HENRY, and L. BASIL

". . . On the 9th instant [October SLOBODKIN. 1878], the sailing-vessel which con- 1953. The size of water masses containing nects us with Key West met water of a plankton blooms. J, Mar, Res, 12(1):141- dark color about midway between here 147, [Dry Tortugas] and there, but saw no dead fish. On her return, on the night "If a phytoplankton population is as- of the 11th, she struck it off Rebecca sumed to be increasing logarithmically Shoals, about 25 miles east of here, in a mass of water surrounded by water and found it extending some 10 miles which is unsuitable for the survival of out in the Gulf. That same night it came the population, it can be shown that down upon us here, and the next morning there is a minimum critical size for the beach and surface of the water, as the water mass below which no increase far as the eye could reach, were cov- in concentration of phytoplankton can

ered with dead fish, . . . Fronn the fact occur, . , . [p, 141.] that almost all the fish that first came ashore were small and of such varieties as frequent shoal water, I infer that the dark water must have been of less "We conclude, therefore, that a popu-

density than the sea. . . . The destruc- lation in a finite region can support

44 for itself against diffusion only if its repro- cases water discoloration extended ductive rate exceeds the leakage, which several miles. He also mentioned an is of the order of magnitude of the ratio outbreak of yellow water caused by of the diffusivity to the square of the Gymnodinium flavum and accompanied extent of the region in the direction of by great luminescence. most rapid diffusion." [p. 146.] KOFOID, CHARLES ATWOOD, and OLIVE KING, GLADYS S. SWEZY. tide in the labora- 1950. Production of red 1921. The free-living unarmored Dino- Inst., 2d Annu. tory. Gulf Carib. Fish. flagellata. Univ. Calif., Mem. 5:1-562. Sess.: 107-109.

Specimens of Gyminodinium breve Gymnodinium sanguineum was cited could not be obtained, but bacteria-free by Okamura as injuring fish in Japan. cultures were made of G. simplex and

marina . Inorganic phos- Plagicampa LACKEY, JAMES B. had no growth- phates and 1956. Known geographic range of Gym- effects. The organisms would promoting nodinium brevis Davis. Quart. J. Fla. in an entirely synthetic medium grow Acad. Sci. 19(1):71, which contains artificial sea water and amino acids added as pure chemicals. Reported obtaining Gymnodinium that dissolved It was recommended breve in two out of four water samples organic nitrogenous matter in Gulf water collected at Trinidad. The samples with be investigated as a clue to blooms. G. breve were taken on March 30 and April 1, 1955, at a temperature of 69 E. KING, JOSEPH to 70.5° F. report on the plankton 1950. A preliminary "According to commercial fishermen, of the west coast of Florida. Quart, there is a yearly fish kill about the Fla. Acad. Sci. 12(2):109- 1 37. J. first of the year, which is not always catastrophic, but might bedue to brevis . Describes, in general, plankton forms So well-known is this that ithas become of the west coast of Florida, especially the basis of a study of areal ocean cur- taken dinoflagellates. Samples were rents as affected by meteorology and to October, 1949. Clarke- from January the flow of the Orinoco River, emptying Bumpus plankton sampler was used for its nutrient rich waters into the Gulf quantitative sampling. Seventy-five of Paria." samples were collected, and listings of plankton taken were given. Although four species of Gymnodinium were LACKEY, JAMES B. listed, G. breve was not included, possi- 1958. Effects of fertilization on receiving bly because all samples were centrifuged waters. Sewage Industr. Wastes at 2,000 r.p.m. for 5 minutes, which 30(11):1411-1415. [Cited from Biol. might destroy the fragile red-tide Abstr.] organism. Adverse effects of fertilization of KOCZY, F. F., M.O. RINKEL, and S. J. receiving waters by sewage or other NISKIN. sources are briefly reviewed. Toxic on the Tortugas 1960. The current patterns effects produced by algal blooms, such Carib. shrimp grounds. Proc. Gulf as the Florida red-tide Gymnodinium Fish. Inst., 12th Annu. Sess.: 112-125. brevis and Prymnesium , which occur in stock- watering ponds, are mentioned. residual cur- A study to determine Adverse esthetic and physical effects rents off the southern west coast of of in such locations as Florida. the Madison, Wis., lakes, and Lake Washington at Seattle, and interferences KOFOID, CHARLES ATWOOD. with treatment of the domestic water re- 1911. Dinoflagellata of the San Diego supply are cited. Extreme diurnal gion. IV. The genus Gonyaulax , with dissolved-oxygen fluctuation may also and a notes on its skeletal nnorphology result from formation of algal blooms. discussion of its generic and specific Author concluded that adverse effects characters. Univ. Calif. Publ. Zool. of admission of mineralized sewage 8(4):187-287. treatment effluents to receiving water Reported great injury to benthic outweigh the possible benefits of stim- living forms. species by Gonyaulax polyedra . In some ulating growth of higher

45 LACKEY, JAMES B., and K. A. CLENDEN- able to assimilate sonne dissolved or- NING. ganic matter, [p, 6.] 1963. A possible fish-killing yellow tide in "The only observed method of repro- California waters. Quart. J. Fla. Acad. duction is by binary fission. Under opti- Sci. 26(3):263-268. mum conditions several divisions may occur in 24 hours, so that bloonns are During the sunnmer of 1961, Gym- easily accounted for. In general, the nodinium flavoim, a pale yellow dino- behavior of this species corresponds to flagellate about 40 microns indiameter, that of dinoflagellates as a whole, but was abundant in San Diego waters, its effects have made it infannous. reaching a maximunn of 1,776 per ml, [p. b.] in one sample. It is suspected of caus- ing small fish kills in Mission Bay. "There have also been rumors of LACKEY, JAMES B., and JACQUELINE A. enteric troubles due to eating shellfish HYNES. from a Red Tide infected area. This 1955. The Florida Gulf coast red tide. point of human illness is emphasized Fla. Eng. Ind. Exp. Sta., Coll. Eng., by La Cossitt (5) in a popular journal, Eng. Progr. Univ. Fla. (9(2):l-23), Bull. but it should be stressed that there has 70, 23 p. been no definite tracing of any human

illness to brevis , and there has been no The report contains a sketch of G. widespread illness when the Tide was breve and the following description of present. The county health officers of the organisnn: the affected area have reported no health "The organism is somewhat discoid troubles fronn it. Dr. Wright, of Sarasota in outline, having a transverse groove County, personally obtained clams dug or girdle, and an indentation or sulcus in a heavily infested area, and bacterio- fronn this girdle to the posterior body logical examination of them by a trained edge, so the body is divided roughly shellfish bacteriologist showed nothing into four quadrants on its ventral sur- unusual. It should be pointed out that face. At the upper end is a small pro- Hornell (loc. cit.) says the people along tuberance or nipplelike projection. The the Indian Malabar coast harvest and sulcus actually extends to the apical eat the fish killed by their Red Tide. end of this nipple, but this can usually No toxic effects like those due to be determined only with the oil imnner- Gonyaulax on the California coast have sion objective, at about 930x. There is been demonstrated. Quite respectable no shell or armor as in some other strings of sheepshead and other fish dinoflagellates. The width is about 30 have been caught in waters containing microns, and the length about 25. There up to 100,000 brevis per liter. These are approximately 22 discoid chromato- were eaten with no untoward effects phores (chlorophyll-containing struc- whatever. Nevertheless, publicity in tures) which appear pale yellow-brown newspapers and nationally circulated in transmitted light. Dense swarnns of magazines concerning actual damage the organisms appear reddish-brown in plus rumored effects has tended to reflected light. The nucleus is typically magnify conditions and probably has large and, as in all dinoflagellate nuclei, further contributed to financial loss in it shows closely packed threads in a the area." [p. 6.] spiral or parallel arrangennent. This is The authors said that estimates of so characteristic, and dinoflagellate fish kills are difficult. They believed nuclei are so resistant to disintegration, that despite the numbers of dead fish as to greatly facilitate identification. In observed, the percentage of kill was

the catenate (chain-forming) Cochlo- low. They stated, ". . . Schools of fin- dinium, formalin used for preservation gerlings have been observed in the very disrupts the cells, but the nuclei still shallowest inshore waters where there stand out and remain connected by was a heavy kill a few hundred yards amorphous remains of the cells. Some- away. The upper part of Tampa and times a few brevis are similarly dis- Hillsborough bays was not invaded by rupted, but the persistent nuclei and the Tide in 1953-54, and at least some chromatophores are diagnostic. The cell large fish were present there. It was contents are clear and finely granular, also observed that while a bad outbreak except that several round refractive with attendant fish nnortality nnight be bodies may be present. No ingested food in progress at one point, a few miles has ever been seen, and the organism away one would find sports fishing and probably lives holophytically, i.e,, like bathing going on as usual, and no scar- a green plant, although it is believed city of mullet for cut bait. At the time

46 of writing, the last localized outbreaks At Sea of the 1953-54 Red Tide are only a few weeks past, yet sports fishing (grouper, SOUTHWEST OF PASS-A-GRILLE, 10 MILES speckled trout, nnackerel, and redfish) has been generally good throughout the September 18, 1953 entire area . . . ." [p. 7.] They gave the following table of fish Huge windrow here, but for several miles an kills: average of 1 dead fish per 10 feet.

COUNTS OF DEAD FISH OFF SANIBEL LIGHT, 6 MILES

March 18, 1954 On Land

Dead fish averaged 1 per 15 feet for an area of LONGBOAT KEY several square miles — perhaps 5x5 miles.

"For purposes of future study, water September 18, 1953 samples containing G. brevis are pre- served by adding 5 ml. of formalin per Fish Counted In Quarter Mile 100 ml. of sample. G. brevis then rounds up and becomes a flattened oval or cir- Yellow Tails 298 Speckled Trout 4 cular disc, usually losing its two flag- lost in a few Whiting U Puffer 1 ella. Color is ordinarily frequently Jack Crevalle 2 Angel Fish i days. Its chromatophores Redfish 3 Porcupine Fish 8 assume a median bandlike position, and indistinct. Pompano I Red Snapper i the girdle and sulcus become Eel 21 Shiner 2 The anterior nipple often can be dis- little practice the Ladyfish 3 Horseshoe Crabs .. 2 tinguished. With very Catfish 11 Unidentified Fish.. 33 species is totally identifiable, even Toadfish 6 after the color has wholly disappeared Triggerfish 2 421 from the chromatophores. [p. 8.] Mullet 11 "In the field, grab samples are taken in shallow water simply by lowering a container from a bridge, dock, or boat, SIESTA KEY and getting an approximate surface sample. For vertical sampling Foerst September 19, 1953 bottles are used, and the depth calcu- lated roughly from meter markings on Fish Counted In 125 Paces (yds.) the line and the angle of the line. This is further correlated with the depth for Grouper 2 Catfish 4 the station as indicated by a Coast and

Whiting 14 Flounder 1 Geodetic Survey map. [p. 8.]

Eel 3 Sheepshead 1 "Preserved samples are sedimented

Porcupine Fish 8 Angel Fish 1 in the dark for two weeks or more, Mullet 14 Yellow Talis 200 when the supernatant may be siphoned

Ladyfish 1 off, and the catch concentrated by cen- Toadfish 7 256 trifuging the final 50 or 100 ml. for five minutes at about 2200 rpm in pointed- end 50 ml. tubes. VENICE JETTIES "Samples to be examined alive are protected from the sun and from any September 19, 1953 sudden increase in temperature. Such living samples should be concentrated Fish Counted in 100 Paces (yds.) and studied within eight or ten hours, also by centrifuging. No tendency to Mullet 418 Other Species... 15 disintegrate has been observed at the speeds used (up to 2300 rpm.), but living organisms will cytolyze within MIDNIGHT PASS 15 or 20 minutes under a cover glass. [p. 8.] December 18, 1953 "Counting is done by a drop method. The catch is concentrated in ratios Fish Counted in Quarter Mile such that 6 drops of catch =100 ml. of raw water. In dense blooms this ratio About 1000 Sheepshead About 8000 Other Species may be doubled or quadrupled, i.e., 12

47 or 24 drops = 100 ml. of raw water. One The area covered was very large, and found taken drop, therefore, equals 16, 8 or 4 ml. of brevis was in samples as raw water, approximately. A 25 mm. far north as Tarpon Springs and as far of square No. 1 cover glass is used, and south as Big Marco Pass, a distance with the usual lOx oculars and lOx or 175 miles, and from land-locked waters 43x objectives in the microscope there such as Lemon Bay to 45 miles off- are 16 or 64 paths--not circular fields, shore. A sample network for an area but paths--across such a cover; that is, this size was not possible, despite the in one drop of catch. If a drop of efforts of the three research organiza- catch =16 ml. of raw water, obviously- tions at work and those of the Gulf Coast one path at lOOx (lOx oculars and lOx Coordinating Committee. In the event objective) equals one ml. of raw water of another outbreak, some such set-up and at 430x one path equals 1/4 ml. to assess the magnitude of the invasion Using the mechanical stage, one counts seems almost a necessity if distribution two paths across each drop on a slide, is to be ascertained, [p. 11.] bisecting the cover and at right angles "At the north and south ends of the to each other. This usually compensates infested area there was a rather sharp for inequality of distribution beneath tapering off of brevis populations. No the cover, and often enables one to fish kills were reported from Tarpon identify much smaller organisms than Springs or Big Marco Pass, although can be identified in a counting chamber. dead fish were reported as abundant off The method is fast and easy, and the Anclote Key on at least two occasions. amount of error is small. The number These may well have been carried there of paths counted depends to some ex- by the northbound current of the eddy tent on the accuracy desired. Common reported by Hela (11). Out in the Gulf, practice has been to count eight paths the most westerly reported occurrences (two paths for each of four drops), of brevis were about 140 miles south- which at 430x usually equals two mis. west of Fort Myers in samples taken by of raw water. G^. brevis is not always the U.S. Coast and Geodetic Survey ship certainly identified at lOOx, but is un- 'Hydrographer.' At the time of this mistakable at 430x. [p. 8.] writing some of the offshore Gulf samples are still awaiting analysis, However, the diminution in numbers

". . . The 1952 outbreak was appar- along the western edge of the shaded ently rather linnited in the area affected, area (Figure 1) indicates that the area which seemed to center around Sanibel of intense growth is fairly well delimited, Island. There is little published infor- [p. 11.] mation on this outbreak, at least at the "Clearwater had a single small fish

present time. . . . [p. 11.] kill along its beaches, with some attend- "The first brevis to appear in 1953 ant discomfort; Naples had a rather were found in samples taken in Lemon larger one. The beaches between these Bay fronn the bridge on August 19, and two points, from Fort Myers Beach to in at the Cortez bridge, the Pinellas county beaches, all had one August 26. The first of these was a or more very heavy fish kills, and the preserved sannple, containing 1 16 brevis inhabitants were subject to a great deal per ml. and the second a living sample, of discomfort. Probably the most con- showing 178 per ml. On the latter date, tinuously affected area was from Casey August 26, there were no brevis at Key to Sanibel, and the heaviest con- Naples or at Piney Point. On September 1 tinuous infestation was in the Boca the organism appeared at the Placida Grande area. [p. 11-12.] Ferry wharf, and on September 3 there were 1,732 per ml. in Big Sarasota Bay,

about three miles north of the bridge, ". . . one cannot detect a discolora- and 600 per ml. in the surf at Golden tion of the water at the surface until Beach on Longboat Key. They were also the number of brevis exceeds 250,000

present as far out as 22 miles off Anna or more per liter. . . . Maria, and on this date dead fish were "Frequently, samples taken a few appearing. It is significant that no dead miles from a rather dense local swarm fish were seen, or at least reported, will show no brevis at all. Examples of until after the populations of brevis had this are easy to cite. On September 18, attained considerable numbers, [p. 11,] 1953, sannples taken just off Pass-a- "Such was the beginning of the 1953- Grille had 2,444,000 per liter, while 54 Red Tide. From the very first it was there were none at John's Pass or in marked by local concentrations rather Sarasota Bay at Cortez. Samples at the than a widespread uniform distribution. latter station contained 83,200 per liter

48 '

small, only small three days later. The whole picture . . . the substrate is very This is one of usually low or absent brevis quantities of charcoal are needed. populations during all of 1953-54, but work is in its preliminary stages, and with sudden erratic, scattered flare-ups there is no field work at present to show of carbon. Experi- to enormous numbers. . . , [p. 13.] the effectiveness ments are underway to show how effec- tively it strips out the growth-promoting "Much attention has been directed matter, how many other organisms are toward recognition of red water from affected, how long its effects last, and boats and planes ... it is frequently to what extent it penetrates (how deep possible to spot patches of bloom in it is effective), [p. 18.] this way. In Peconic Bay it has been "Carbon has three other characteris- possible to spot blood- red concentra- tics which recommend it. It is rela- tions of a related dinoflagellate, Cochlo- tively cheap, and the supply is inex- dinium as far as a half-mile away from haustible and readily available. It is a small cabin cruiser. However, no not in itself toxic; it will not harm fish. such vivid discolorations have been There is no reason why it should affect noted in hunting for brevis either by organisms which have no organic re- boat or by air, and red water has fre- quirements. Finally it has been shown quently been all but impossible to locate, by Block (17) that it will adsorb the or has been a slightly brown discolora- water borne toxin of brevis . The effect tion rather than red. . . . any water of this would be to cleanse water located in such a manner must be ex- already poisoned. Clearly charcoal and amined microscopically. . . . [p. 13.] activated carbon need a very careful investigation, [p. 18.] "Biophysics has demonstrated many ways, principally through wave action, microorganisms. ". . . Considering the tendency of phy- of affecting living toplankton to accumulate in the topmost Here again the work at the University few inches in lakes, ponds and even of Florida is just beginning. It has streams as a direct response to light, already been shown, however, that there it is evident that brevis presents no is no useful killing action in a high- such clearcut response. ... a trend to frequency radio field. Cultures of five accumulate in deep water is sometimes organisms, including brevis , were un- evident and will be more closely in- affected in glass containers in such a vestigated in future work. [p. 14-15.] field. ... [p. 18.] phosphorus as ". . . Since much of our idea of the The authors mentioned distribution of brevis has been based a possible cause of red tide and gave until now either on surface samples or the following table of phosphorus values. (perhaps) on dead fish, these observa- tions suggest that currently accepted ppm distribution patterns, or areas of in- 2-19-53 Peace River, Arcadia, Odum 0.850 festation, be accepted tentatively, for 2-19-53 Peace River, Punta Gorda, Odum. 0.240 depths greater than ten feet. [p. 15.] 3-30-53 Placlda (Peace Estuary), Odum.... 0.028 4- 1-53 EI Jobean (Peace Estuary), Odum. 0.144 4-17-53 Bokeelia (Peace Estuary), Odum.. 0.024 3- 7-54 Peace River, Punta Ckjrda, Odum "... In the light of past experiences and Hynes 0.670 with algicides, it seems probable that 3-18-54 St. James Point (Peace Estuary), one of rather high specificity and very Odum and Hynes 0.040 high killing power can be developed for 4- 3-53 Klssimmee River above L. Okee- 0.018 brevis . . . . control of brevis in the chobee, Odum shallow inshore areas along the coast 2-19-53 Lake Okeechobee, Moore Haven, by the use of plane-dispersed algicides Odum 0.039 must not be discounted, until very 2-19-53 Caloosahatchee Estuary, Sanibel, thoroughly evaluated, [p. 18.] Odum 0.009 ' Gymnodinium brevis requires small 2-19-53 Caloosahatchee Estuary, Punta quantities of organic substances in its Rassa, Odum 0.060 nutrition. Wilson (13), in his culture 3-30-53 Caloosahatchee Estuary, Ft. medium, includes vitamins and soil ex- Myers Beach, Odum 0.042 tract. These are present to some extent 3- 6-54 Caloosahatchee Estuary, Ft. in Gulf water, and Odum, Hynes and Myers, Odum and Hynes 0.094 Slater (16), have shown that charcoal 3- 6-54 60 miles off Naples, Odum 0.001 may effectively remove them by ad- 3- 6-54 70 miles off Naples, Odum 0.042 sorption. Since the organic portion of 3- 6-54 80 miles off Naples, Odum 0.001

49 ppm Davis [personal communication] says 3- 6-54 110 miles off Naples, Odum 0.001 exposure of mangrove peat by storms 4- 1-53 Hillsborough Bay, Odum 0.410 might be a factor. This peat contains 4- 7-53 Piney Point, Tampa Bay, Odum.... 0.120 H2S when first exposed, and has been 4- 7-53 Pinellas Point, Tampa Bay, Odum 0.108 accumulating for at least 5,000 years. 4- 6-53 Ballast Point, Tampa Bay, Odum.. 0.580 Davis says it is four feet thick at Naples, 4- 6-53 Gandy Bridge, W. end Tampa Bay, but presumably it is equally thick on Odum 0.240 the Atlantic coast of Florida, [p. 21.] 3- 7-54 Tampa Bay, Odum and Hynes 0.410 "Organic acids (humic, tannic) are abundant in runoff from the West Coast, Their reference in the table to Odum and [viewed] from the air, the brown is apparently to a personal communica- discoloration as this water comes out ." tion from Howard T. Odum, and their of the passes is spectacular. . . [p. reference to Odum and Hynes is appar- 21.] ently Odum, Hynes, and Slater which Collier (1953b) "showed that titanium they listed as a report in preparation and zirconium were significantly present entitled "Suggestion concerning the use in the brevis area, but of themselves of charcoal for control of 'red tide' they have failed to be a stimulant in blooms. laboratory cultures. . , ." [p. 21.] The authors disagreed with the con- clusion of Chew (1953) that phosphorus "All of the samples in the list above, is lower in river water than Gulf water. except the last, indicate a progressive The authors discussed the suggestion diminution of phosphorus as the water of Graham (1954) concerning the pos- ." travels into the Gulf. . . [p. 19.] sible role of Skujaella ( Trichodesmium ) The authors pointed out that to obtain in causing blooms by concentrating the phosphorus values given by Ketchum phosphorus, but concluded that, "Neither and Keen for a 3-foot surface layer in the literature nor observations of this a depth of 30 feet, all of the phosphorus laboratory over the past few years have would be concentrated by the organism borne out the idea of an extensive algal in this surface layer. If the whole column bloom of blue-greens preceding Red was this rich in phosphorus, 17,000 Tide outbreaks." [p. 22.] pounds of pure phosphorus per square Meteorological and current effects mile would be required. They then sum- "certainly come into play in distributing marized by stating, "Actually brevis the land -contributed nutrients and in has been found blooming in water prac- dispersing the organisms, and meteoro- tically lacking in detectable phosphorus logical effects are seen when heavy (June 18, 1954, Center of Lemon Bay, weather breaks up a heavy brevis con- Tables IV and VII: phosphorus 0.000 centration, [p. 22.] ppm, brevis 4,814,000 per liter) and in "... Odum has measured the light water containing abundant phos- intensity in varying concentrations of phorus. . , ." [p. 20.] brevis . and while at the surface inten- The authors discussed nitrogen as a sities may be very high, the organisms possible factor in blooms. They stated have been found concentrated at depths that Odum (personal communication) where the light intensity was greatly failed to find any nitrate nitrogen in a reduced. These and other considera- series of about 15 field samples in tions would seem to rule out light as a brevis water, Oct. 6, 1953. They quote single decisive factor." [p. 22.] him as stating "the suggestion is thus nnade that Gymnodinium [ brevis ] is not LACKEY, JAMES B., and C. N. SAWYER. a nitrate user." [p. 20.] 1945. Madison lakes survey. II. Biological aspects. In Investigations of the odor nuisance occurring in the Madison lakes "Except for changes in Gulf water due particularly Monona, Waubesa, Kegonsa to dilution with land run- off, there from July 1943 to July 1944, p. 67-72. seems little else which is peculiar to Rep. Gov.'s Comm., Wis. [Cited from the brevis area. Ground water in the Lackey and Hynes, 1955. Non vidi.] general area is frequently highly charged LaCOSSITT, HENRY. with H2S. . . However, the ground-water 1954. The truth about Florida's Red Tide, springs in the Gulf flow constantly, and Sat. Eve. Post 227(7):28ff. [Cited from there are perhaps as many on the Lackey and Hynes, 1955.] Atlantic side, where brevis has not been found. There is no reason to sus- Lackey and Hynes (1955) said con- pect that the flow is excessive during cerning this article, "This point of intermittent brevis outbreaks, John H. human illness [enteric troubles from

50 .

." 175- eating shellfish from a red tide infected this explanation doubtful. . . [p. area] is emphasized by La Cossitt (5) 176.] in a popular journal." LEBOUR, MARIE V. LANGMUIR, IRVING. 1925. The dinoflagellates of northern seas. 1938. Surface motion of water induced by Mar. Biol. Lab., Plymouth, England, wind. Science 87(2250):119-123. [Cited 172 p. Ryther, and Pomeroy et al., by 1955, LEES, G. M. 1956.] 1937. "Black Sea" conditions in the Arabian Discusses the accumulation of float- Sea. Amer. Ass. Petrol, Geol., Bull. ing objects in streaks parallel with the 21(12):1579-1582. prevailing wind by wind-driven convec- LEWIS, GEORGE J., JR., and NORRIS W. tion cells. RAKES TRAW. 1955. Carbohydrate in sea water, J. Mar, LASKER, REUBEN, and F. G. WALTON SMITH. Res. 14(3):253-258. 1954. Red tide. In Paul S. Galtsoff (coordi- nator), Gulf of Mexico: its origin, Gives anthrone method of determining waters, and marine life, p. 173-176. carbohydrates Fish Wildl. Serv., Fish. Bull. 55. LONG, E. JOHN. This paper reviews red-tide out- 1953. The red tide hits and runs. Nature might cause breaks and factors which Mag. 46(3):125-128. [Cited from Biol. of Gymnodiniunn brevis . overblooming Abstr.] The authors discounted the theory that blooming G^. brevis is caused by up- The occurrence of the "red tide"

welling of nutrient- rich water along dinoflagellate, Gymnodinium brevis , is Florida's west coast. discussed. Water samples taken during One possible cause given was in- red tide have excess of N and P. Some creased phosphorus concentration, relation is thought to exist between the either by accunnulation by G^. brevis heavy discharges of fresh water from itself or by convergence of phosphorus- the rivers of southwest Florida and the rich water masses. occurrence of the red tide. It is also theorized that "The con- centration of nutrients may have been LUCAS, C. E. the result of the lateral or vertical 1947. The ecological effects of external migration of some organism or or- metabolites. Biol. Rev. 22(3):270-295. ." ganisms other than Gymnodinium . . . [Cited from Ryther, 1955.] [p. 175.] LUCAS, C. E. personal communication, George In a 1949. External metabolites and ecological of University theo- L. Clarke, Harvard adaptation. Symp. Soc. Exp. Biol. III. rized that "vertical migration linked Selective toxicity and antibiotics, p. concentra- with a subsequent horizontal 336-356, Academic Press, Inc., New of water tion due to convergences York. [Cited from Ryther, 1955.] masses may be the complete solution." In 1950, phosphate deposits covering LUND, E. J. a 25-mile area were discovered off 1935. Some facts relating to the occur- of dead and dying fish on the Tannpa, Fla. ". . . The coincidence of rences these deposits with the area in which Texas coast during June, July, and red tide outbreaks have occurred sug- August 1935. Tex. Game Fish Oyster gests that this may be a partial explana- Comm., Annu. Rep. for the fiscal year tion of phosphorus availability. 1934-35:47-50. "The problem presented is to account Gives observations on dead fish salts for periodic releases of nutrient washed ashore in the vicinity of Port from these bottom deposits. Possible Aransas from June 30 to August 13. shifts due to cata- explanations include Most of the fish were in an advanced or clysmic upsets in the ocean bottom state of decay when they washed ashore. of bottom simple mechanical shifting "The escape of irritating 'gas', dis- bottom cur- muds due to strong solved in the sea water was facilitated ." rents. . . [p. 175.] by prevailing winds blowing on shore. suggested as a River drainage was Appearance of 'gas' was always asso- source of mineral deposits; however, ciated with the simultaneous or imme- ". of places like . . The remoteness diately previous appearance of dead Key West and Cape Sable from river fish. ... [p. 48.] drainage areas where rich phosphate deposits are known to exist renders

51 ". . . an examination of the Weather however, they may be seen to be Bureau reports on Texas for 1934-5 clustered thickly about a cell of the shows that rainfall and flood levels of Gymnodinium . This gelatinous envelope the rivers were at unusual naaxima nriay well be a factor of importance in during May and June, 1935. A maximunn holding the organisnns together, once of the resulting outflow of water of very they are massed by a favorable com- low salinity throughthe passes occurred bination of light, water temperature, during the last week in June and there- and tidal currents." [p. 223.] fore just preceding the first appearance MARVIN, KENNETH T, of dead fish on Padre Island. . . ." [p. 1955. Oceanographic observations in west 49.] coast Florida waters, 1949-52. Fish MARGALEF, RAMON. Wildl, Serv., Spec. Sci. Rep. Fish, 149, 1963. Succession in marine populations. i + 32 p. (editor), Advancing fron- In Raghu Vira This report presents raw hydro- plant sciences, vol. 137- tiers of 2, p. graphic data collected by the Fish and 188. Institute for the Advancement of Wildlife Service from May 1949 through Sciences and Culture, New Delhi 16, July 1951. Data do not extend into 1952 India. as the title of the paper implies. Data MARTIN, G. W., and THURLOW C. NELSON. were collected by the vessel Pompano 1929. Swarming of dinoflagellates in Dela- from 10 stations (1 in Charlotte Harbor ware Bay, New Jersey. Bot. Gaz. 88(2): and 9 offshore to as far as 120 miles). 218-224. Two other stations were in the Peace and Caloosahatchee Rivers. Station data They described discolored water oc- include temperature, salinity, oxygen, curring in late summer of 1928 in inorganic and total phosphorus, pH, Delaware Bay, N.J. It extended from and nitrites. the shore to about 7 m. out, where the MARVIN, KENNETH T. depth was 2 m. The chief organisnn 1958. Copper ore experin-ients for red tide was Annphidinium fus ifor me . Oysters control. Jn Annual report of the Gulf exposed during low tide were examined Fishery Investigations for the year after being covered half a meter and ending June 30, 1958, U.S. Fish Wildl. were found to be full of the organisms. Serv., p. 78-85, Some patches of "red" water con-

tained Gymnodiniunn . They found when Most of this material is discussed they were killed with a strong solution in Marvin et al. (1961).

of iodine in potassium iodide and ex- ". . .the source of copper would have amined microscopically that each cell to be capable of releasing a relatively was surrounded by a gelatinous envelope small amount of copper into solution as thick as the diameter of the cell it- over a long period of time. , . .The self, so that the water in which they particular ore under consideration is occurred must have been a nearly con- a sulphide ore containing roughly 1 per- tinuous mass of soft jelly. The envelope cent copper, 3.5 percent iron and 6 per- is also visible, but less apparent, if the cent sulphur. organisms are killed in a saturated "Preliminary investigations have solution of bichloride of mercury. The ,shown that the ore in a closed system envelope is less regularly seen in ma- (4 pounds of ore in 120 gallons of sea terial killed in osnnic acid and then trans- water) will liberate a maximum of ferred to chrom-acetic solution, and about 1.5 parts per n^illion of copper never in any of the other reagents used. in about a day's time. This then tapers

". . .Gelatinous envelopes are com- off to about 0.9 part per million; the mon among dinoflagellates when en- excess copper is presumably precip- cysted, but not when active. The cells itated as the rather insoluble com- ." referred to in this connection were pound, basic copper carbonate. . . actively motile. A similar envelope [p. 78-79.] has been noted occasionally surround- MARVIN, KENNETH T. ing other species of naked dinoflagel- 1959, Testing copper ore for red tide con- lates in the active condition, but only trol. In Galveston Biological Laboratory the iodine or bichloride when killed by fishery research for the year ending methods. In red water plankton in which June 30, 1959, p. 86-94. U.S. Fish Amphidinium fusiforme is the dominant Wildl, Serv., Circ. 62. species, the Amphidiniunn cells tend to cling together in clumps, but no gelati- "Field tests have been conducted to nous envelope can be demonstrated. In determine the effects of copper sulphate many of the clumps (although not in all). on G. breve when applied as crystals

52 —

and also as a concentrated solution. These tests were conducted during red tide outbreaks on the west coast of Florida in 1952, 1953, 1954, and 1957. Copper sulphate dissolves rapidly in sea water supersaturating the ambient water with copper. The resulting con- centration is many times that of the required lethal level as determined in laboratory toxicity experinnents, but this high concentration does not last long. The bulk of the copper precip- itates as the basic carbonate and is dispersed by tides and currents and the copper concentration is reduced to an ineffective level." [p. 87.] The report gives details of the leach- ing rate of copper ore and of the copper ore experiments. For the re- sults see Marvin et al. (1961).

MARVIN, KENNETH T, 1960. Chemical control experiments, fii Galveston Biological Laboratoryfishery research for the year ending June 30, 1960, p. 43-45. U.S. Fish Wildl. Serv., Circ. 92.

The need is pointed out for a chemical toxic to G^. breve at low concentrations, selective as to the organism it affects, and without the precipitating character- istics of copper. Results of 16 replicate experiments at six concentrations of CUSO4. 5 H2O indicated that the critical concentration of copper for 100-percent mortality of G. breve lies between 0.39 and 0.80 ug.at./l. The screening of 4,000 compounds revealed the following numbers toxic to G. breve at five concentrations shown below:

Conpounds containing The compounds toxic at 0.04 p. p.m. "ciguatera" fish poison rather than the are being tested against other marine usual paralytic shellfish type. Samples organisms to determine their specificity from all five Gulf States in December for the red-tide organism. 1962, were negative except for oysters of Sarasota and Charlotte Counties. MATODA, S. 1944. Sea and plankton. Kawade Shobo Publ., McFARREN, E. F., M, L. SCHAFER, J. E. FishandCobb, 1954.] 235 p. [Cited from CAMPBELL, K. H. LEWIS, E. T, JENSEN, According to Fish and Cobb, Matoda and E. J, SCHANTZ. said observations by K. Nakazawa in- 1957. Public health significance of paralytic dicated that early larvae of mollusks shellfish poison: a review of literature and crustaceans, which breathe through and unpublished research. Proc. Nat, the body surface, survive in discolored Shellfish. Ass. 47:114-141. water, while older forms with special- ized respiratory gills are killed when The significant feature of this paper these organs become completely is failure to mention either G. breve covered with the disintegrating bodies or oysters in connection with shellfish of dinoflagellates. These and similar poisoning. See McFarren et al. (1960). observations by K. Oda on oysters in Gokasho Bay when Gymnodinium McFARREN, EARL F., MARY L. SCHAFER, mikimoti Miyaki and Kon-iinani swarmed JEPTHA E. CAMPBELL, KEITH H. LEWIS, in 1934, led to the conclusion that the EUGENE T. JENSEN, and EDWARD J. detrimental effect is of a physical SCHANTZ. rather than chemical nature. i960. Public health significance of paralytic shellfish poison. Advances Food Res. MAY, BILLIE Z. 10:135-179. 1964. A collection of data in reference to red tide outbreaks during 1963. 6. En- In this report it is stated that the vironmental chemistry: January through only known case of Fla. Bd. Conserv., Mar. May, 1963. from eating oysters occurred inBaynes Lab., 108-121. p. Sound (western side of the Gulf of For 18 stations in Tampa Bay and Georgia), British Columbia, from eat- adjacent waters in January through ing cultivated Japanese oysters,

May, 1 963, author gives salinity, oxygen, Crassostrea gigas .

calcium, total phosphate, surface chlo- ". . .The appearance of the poison in rophyll A, and ultra-violet absorption the oysters was accompanied by a values (given as optical density on a much higher level of toxicity in butter Beckman DU spectrophotometer with clams in adjacent areas, while little

a 1 cm. cell; measured 220 m^ ). Samples neck clams, razor clams, and cockles were filtered before measurements with exhibited a relatively lower level of a millipore HA filter of 0.8 pore size. toxicity. In this instance, the oysters The author ascribed the variations in appeared to rid themselves of the poison the optical-density readings to the pres- more rapidly than other bivalves after ence of dissolved organics, citing Arm- the ingestion of the toxic plankton had strong and Boalch (1961). The optical- ceased." [p. 145.] density readings are fairly well correlated with salinity. For nine of McKEE, JACK EDWARD, and HAROLD W. the K stations taken each month from WOLF (EDITORS). February through May a correlation 1963. Water quality criteria. 2d ed. The of -0.75 indicates much higher absorp- Resources Agency Calif. State Water tion of light as the sea water becomes Quality Control Board, Publ. 3-A, diluted with river water. Plotting of 548 p. the optical- density readings against chlorophyll A for the same stations The section on marine flagellates shows also an increase in optical den- (p. 323-324) gives various summary sity as chlorophyll A increases. statements and sources from whichthey were derived. The statements, however, McFARREN, EARL F. have not been updated; so theories put 1963. illness produced by eating Human forth a number of years ago which Sarasota and Charlotte County, Florida, seenned reasonable in the light of exist- oysters, I. Analysis of sannples. Robert ing knowledge, but which have been A. Taft Sanitary Eng. Center, Public superseded by later work, are given as Health Service, p. 1-4. facts. For instance, it is stated that Assays on mice suggested that the sulfides are essential for mass develop- poison from the oysters resembled ment of the red-tide organism, giving

54 as authority an anonymous "fill in" note galbana was satisfied by inorganic on the bottom, of a page of the Progres- phosphate, commercial glycerophos- sive Fish-Culturist for July 1953. phate, yeast adenylic acid, cytidylic acid, monoethyl phosphate, and ribo- Mclaughlin, john j. a. flavin monophosphate. 1956. The physiology and nutritional re- "Laboratory cultures in defined media quirements of some chrysomonads. of the isolates of P. parvum were Ph.D. Thesis, New York Univ., 65 p. toxigenic to Lebistes and Gambusia . [Reference from Ray and Wilson, 1957, Culture fluids from alkaline media were p. 493. Non vidi.] more toxic than those from acid media, as previously noted in Israel. Ray and Wilson (1957) stated, "Culture media suitable for produc- "McLaughlin working with the sameor- tion of large quantities of these or- s ganisms were developed." [p. 75.] ganisnn | P r ymne ium parvum ] reported that cultures grown in an alkaline medium were more toxic than those McLaughlin, john j. a., and luigi grown in an acid medium. P. parvum PROVASOLI. cultures (grown in alkaline nnedia), 1957. Nutrition requirements and toxicity rendered nontoxic by lowering the pH of two marine Amphidinium . J. to 6.0, regain their toxicity when made Protozool. 4 (suppl.):7. alkaline (Shilo and Aschner, 1953; McLaughlin)." Amphidinium klebsii (?) and A.

rhyncocephalum require vitamins Bj2 . Mclaughlin, j. j, a. thiamine, and biotin. 1958. Euryhaline chrysomonads: Nutrition

and toxigenesis in Prymnesium parvum , MEAD, A. D. with notes on Isochrysis galbana and 1898. Peridinium and the 'red water' in

Monochrysis lutheri . J. Protozool. 5( 1 ): Narragansett Bay. Science, n. s. 8(203): 75-81. 707-709; also in Rep. U.S. Comm. Fish Fish. 1899, p. CXLIL "The nutritional requirennents of 3 isolates of Prymnesium parvum (2 Is- MENON, M. a. S.

raeli, 1 Scottish) included vitamin Bj2 1945. Observations on the seasonal dis- and thiamine. For comparison, 2 other tribution of the plankton of the brackish chrysomonads were studied: Trivandrum coast. Proc. Indian Acad. Monochrysis lutheri isolated by Droop Sci. B22:31-62. [Cited by Ryther, 1955.] in Scotland Isochrysis galbana puri- and Blooms of Gymnodinium sp. occur fied by McLaughlin from a culture ob- annually during June to September along the laboratory. tained from Plymouth the Trivandrum Coast of India, following parvunn "The isolates of Prymnesium rains and diatom maxima. and Isochrysis galbana had a molecular Bj2 specificity like Ochromonas MITSUKURI, K. malhamensis : no response to Factor B, 1905. The cultivation of marine and fresh- pseudovitamin Bj2 , Factor A or Factor water animals in Japan. U.S. Bur. responded H. M^. lutheri , in contrast, Fish., Bull. 24:259-289. to pseudovitanain B i2> Factor H, and Factor A. "As in all culture enterprises, there "Thiannine was essential; 1.0 ug. % are many enennies of the pearl oyster, difficulties in the allowed full growth of P. parvum . The as well as unexpected NaCl concentration foF good growth way of its culture. Octopus , Codium, was 0.3-5.0%; growth was possible from Clione (sponges), all sometimes play a 6-12%. Dark growth was not achieved. sad havoc among the mollusks, but the "Ammonia, as suggested from its use most dreaded enemy of all is the 'red 'red tide.' This is an im- in suppressing outbreaks of P. parvum , current' or was sharply inhibitory, less so at high mense accumulation of a Dinoflagellata, concentrations of NaCl or at acid pH. Gonyaulax, causing discoloration of the "Nitrate, annmonia, arginine, aspara- sea water, and, in some way not well gine, naethionine, histidine, alanine, gly- accounted for, causing in its wake an cine, serine, proline, leucine, isoleu- immense destruction of marine orga- cine, tyrosine, aspartic and glutamic nisms, large and small." [p. 284.] acids, acetylurea, and creatine served as nitrogen sources in both acid and MIYAJIMA, M. alkaline media. 1934. La question de "I'eau rouge" un peril "The phosphate requirement of P. pour les huitres perlieres. Bull. Soc. parvum and M. lutheri and Isochrysis Gen. Agr. Peche 41:97-110. [Non vidi.]

55 MOE, MARTIN A. NISHIKAWA, T.

1964. A collection of data in reference to 1901. Gonyaulax ( Polygamma ) and the dis- red-tide outbreaks during 1963. 7. A colored water in the Bay of Agu. Annot. note on a red tide fish kill in Tampa Zool. Jap., Part I, IV: 31-34. Bay, Florida, during April 1963. Fla, Bd. Conserv., Mar. Lab., p. 122-125. An outbreak of red water by Gonyaulax polygamma did no dannage; earlier out- Identified 52 species of fish killed in breaks in some localities were reported the outbreak. to be highly destructive to pearl oysters, fishes, and crustaceans, MONTAGNE, C. 1844. Sur le phe'nomene de la coloration des NORDLI, E. eaux de la mer rouge. Ann. Sci. Nat,, 1953. Salinity and temperature as control- ser. 3, Bot. 2:332-362. [Non vidi.] ling factors for distribution and n-iass occurrence of Ceratia. Blyttia 11:16- MOORE, M. A, 18. [Cited from Ryther, 1955.] 1882. Fish mortality in the Gulf of Mexico, Proc. U.S. Nat. Mus. 4:125-126. NUMANN, WILHELM. 1957. Natiirliche und kiinstliche "red water" "About two years ago [1878] certain mit anschliessenden Fischsterben im portions of our Gulf waters became Meer. (Natural and artificial "redwater" poisoned in some way that caused the with associated fish mortalities in the death of all the fish that came in con- sea). Arch. Fischereiwiss. 8(3):204- tact with it. Whenever a smack with a 209. [Summarized from a translation full fare, i.e., a full cargo of fine healthy by Alexander Dragovich.] fish in her well, sailed into this poisoned water every fish would die, and they He personally observed in the would have to be thrown away. Bodensee how whitefish avoided the "This state of affairs has occurred areas having heavy diatonn concentra- again [1880]; the waters of some por- tions. tions of the Gulf becoming so noxious Author observed an outbreak off the as to kill the fish. The poison seems to coast of Angola during August 1951. He be confined to certain localities and stated that the nutrient-rich zone ex- currents for the time being, as some- tends from Angola to Lobito-Benguela, times this state of affairs is observed where the Benguela current deflects to more marked at one place and some- the west. At the end of July and the times at another. However, there seems beginning of August he observed an to be more of it about the mouth of explosionlike outburst of phytoplankton Charlotte Harbor and off Punta Rassa a little north of the mouth of the Congo. than elsewhere." [p. 125.] As he traveled south, yellowish-brown and red masses of plankton occurred MUNK, WALTER H., and GORDON A. RILEY. for 600 km, to Lobito, north of the 1952. Absorption of nutrients by aquatic nutrient-rich coastal waters. The plank-

plants. J. Mar. Res. 1 1(2):2I 5-240. ton masses were never observed on [Cited by Ryther, 1955.] the surface farther than 10 miles off- shore. A deep red was seldom ob- NELSON, THURLOW C. served; instead, a brownish. yellow was 1948. Red oysters. N.J. Fisherman (Nov.) much more frequently noted. [Cited from Pomeroy et al., 1956. Non During the first half of September vidi.] 1951, water in the Bay of Luanda was blood- red, and enormous quantities of Red oysters, exuding blood-red dead fish covered the surface. He took liquor, were associated with dinoflagel- plankton sannples 5 miles off the coast late blooms in Delaware Bay. of Luanda on July 27, at the beginning of the outbreak. In individual samples NIGHTINGALE, H. W. (analyzed by De Sousa e Silva, 1953) 1936. Red water organisms. Their occur- 69 to 92 percent was Exuviella baltica rence and influence upon marine aquatic Lohmann. Prorocentrum micans Ehren- animals, with special reference to shell- berg and Prorocentrum sp. constituted fish in waters of the Pacific Coast. The up to 10 percent, and Peridinium sp. Argus Press, Seattle, Wash. 24 p. [Non up to 1 3 percent. vidi.] He described an outbreak of red tide in the Bay of Izmir (Agean Sea) caused

Reviewed red-water worldwide occur- by a species of Gymnodinium . The rences since 1871. water was coffee brown to yellowish

56 or greenish. He speculates on the cause 10 small acid streams draining phos- p.m.; and of the fish mortality, ascribing it par- phatic formations as 0.413 p. for- tially to suffocation and partially to 7 streams not draining phosphatic toxic substances. mations but receiving sewage the phos- A translation of his remarks on both phorus as 0.836 p. p.m. outbreaks follows: "The outburst of red water never occurs in the open sea, ODUM, HOWARD T., J. B. LACKEY, but always in bays and coastal waters JACQUELINE HYNES, and NELSON MAR- which are rich in nutrients. According SHALL. to all reports, outbreaks of red water 1955. Some red tide characteristics during nnost obviously occur in coastal areas 1952-1954. Bull, Mar. Sci. Gulf Carib. a short time after precipitation. The 5(4):247-258. bloom of Exuviella occurred in the A report on miscellaneous red-tide after heavy coastal area of Angola data (total nitrogen, phosphorus, and The rainfall in the hills of Binnenland. counts of G. breve --45 determinations Congo and Cuanza Rivers brought much with both N and P) from coastal waters. water which was spread throughout the Summary table does not give range of coastal waters. surface layers of values in the five localities sampled. "It also appears- -and this is our Other data are mentioned (chlorophyll, -that a outburst final conclusion- mass light measurements, and B.O.D.) but fresh- of phytoplankton occurs when not presented, although some results water growth-promoting substances are discussed. G. breve counts were elennents, enzynnes, or other (trace made from samples preserved in 5 per- biologically active substances) reach cent formalin and centrifuged. The data the sea. Due to the presence of a nec- are rather meager for reliability. Odum nutrients in the essary quantity of said Olson (1953) showed that the fresh sea, a pre-condition for the outburst of water entering Tampa Bay takes 2 to 3 in the exists already. plankton sea years to accumulate a volume equal to only Accordingly, these outbursts occur that of the bay and, thus, to complete a near the coast." turnover. The increasing multiple pollution of ODUM, HOWARD T. the shallow coastal estuaries is given in wa- 1953. Dissolved phosphorus Florida as possibly the cause of a higher inci- State and Fla. ters. Fla. Bd. Conserv., dence of red tide than might otherwise Geol. Surv., Rep. Invest. 9(l):l-40. occur. A footnote mentions that Odum and deposits phosphate "The extensive of Hynes, at the 1954 meeting of the Amer- lead to unusually high rock in Florida ican Society of Limnology and Oceanog- contents in the dissolved phosphorus raphy in Gainesville, Fla., suggested drain these streams and lakes which the large-scale dusting of charcoal . quantities of dis- areas. . . Additional as a possible means of absorbing re- solved phosphorus are being added by quired vitamins and, thus, modifying areas, sewage and industry in some the nature of offshore blooms. although little recognition has been made of the possibly large biological OLSON, F. C. W. amounts of effects that relatively small 1953. Tampa Bay studies. Fla. State Univ., have on those added phosphorus can Oceanogr. Inst., Rep. 1:1-27. areas which are not receiving drainage from phosphate areas. The moderately Fresh waters entering Tampa Bay content of basic springs low phosphorus take 2 to 3 years to accumulate a vol- surface streams in contrast to acid ume equal to that of the Bay. suggests a controlling role of pH in Florida." phosphorus solubility in [p. 1.] OTTERSTR0M, C. V., and E. STEEMANN This report contains several maps NIELSEN. phosphate-bearing forma- showing the 1939. Two cases of extensive nnortality in dissolved tions and the concentrations of fishes caused by the flagellate, phosphorus in Florida waters. Separate Prymnesium parvum , Carter. Rep, River, figures are shown for the Peace Danish Biol. Sta. 44:5-24. the Tampa Bay, the South Florida, and of a the St. Johns River areas. "The effect of the poisoning is He stressed the effect of sewage in permanent nature, the conveyance of an greatly increasing the phosphorus con- apparently unhurt fish to pure water tent of lakes and streams. Thus, he does not by far in all cases save the gave the phosphorus for 18 small un- fish." The flagellate itself is not poi- polluted streams as 0.019 p. p.m.; for sonous; the damages must be caused

57 .

by some matter contained in the water all linriited to the surface layer of the and most likely originating from the water, even in the absence of a sig- metabolism of the flagellate. nificant density gradient. In addition to being concentrated at the surface, PAULSEN, OVE. the blooms were typically in the form 1934. Red "water bloom' in Iceland seas. of elongated slicks with abrupt margins.

Nature 1 34(3399):974. This was also true in the 1952

bloom. . . . PEARSALL, W. H. "No mortality of marine organisms 1932. Phytoplankton in the English lakes. was associated with the blooms. How- II. The composition of the phytoplank- ever, a few days after the appearance ton in relation to dissolved substances. of the 1952 bloom local oyster packing J. Ecol. 20(2):241-262. [Cited from companies began to complain of 'red Ryther, 1955.] oysters'. The red material was con- centrated in the digestive glands of the PETERS,_N. oysters. After several days in cold 1929. Uber Orts- und Geisselbewegund bei storage the liquor of the shucked oysters marinen Dinoflagellaten. Arch. became blood-red. Several weeks later, Protistenk. 67(2/3):291 - 321 oysters in nearly all parts of Delaware Bay had developed dark red digestive Ponaeroy et al. (1956) cited Peters glands. They no longer exuded red liquor (1929) and Hasle (1950, 1954) as sug- after being shucked, and the wet volume gesting that some species of dinoflagel- of meats per bushel of oysters in- lates can oppose successfully a vertical creased from 4 or 5 pints per bushel current of about 1 cm. /sec. to 8 " [p. 56.] The authors described several blooms PHELPS, EARLE B., and DAVID E. BARRY. of Gymnodinium sp. in the headwaters 1950. Stream sanitation in Florida. Univ. of Duplin River, a wholly tidal stream, Fla., Eng. Ind. Exp. Sta., Bull. 34:1-56. with no vertical or horizontal salinity gradient. At the time of one bloom Gives amounts of sewage from St. (April 21, 1955) it was noted that the Petersburg and Tampa. bloom disappeared at night and in the early morning on 4 successive days, PIERCE, H. D, reappearing each day at about 0900. As 1883. 53-- The spawning of bluefish--an the bloom was concentrated in several opinion of the cause of mortality of patches near the downwind shore, they fish in the Gulf of Mexico. U.S. Fish released widely spaced current drags Comm., Bull. 3:332. 10 cm. in depth, ballasted to float in the upper 10 cm. of water. These drags This reference, cited by others, has demonstrated convergence of surface no bearing on red tide. water by all drifting and remaining close together in the center of a dense PINTNER, I. J., and L. PROVASOLI. concentration of dinoflagellates. 1963. Nutritional characteristics of some The organisms were phototactic; by chrysomonads. fa. Carl H. Oppenheimer moving the light source it was found (editor). Symposium on marine micro- they moved between 0.3 and 0.8 cm./ biology, ch. 11, p. 114-121. Charles C. sec. Thomas, Springfield, 111. "The observation that blooms occur only during light winds suggests that POMEROY, LAWRENCE R,, HAROLD H. HAS- at wind velocities of Beaufort 3 or KIN, and ROBERT A. RAGOTZKIE. more the speed of the current vortices 1956. Observations on dinoflagellate exceeds the swimming speed of the

blooms. Limnol. Oceanogr. l(l):54-60. dinoflagellates. . . ." [p. 58.] They cite Peters (1929) and Hasle (1950,1954) Discusses dinoflagellate blooms in as confirming the suggestion that some Delaware Bay and in tidal creeks on species of dinoflagellates can oppose Sapelo Island, Ga. The Delaware Bay successfully a vertical current of blooms were caused by a mixture 1 cm./sec. They then say that of Annphidinium fusiforme and "Langmuir's (1938) measurements sug-

Gymnodinium splendens . ". . .AH the gests that this vertical velocity is ex- Delaware Bay blooms occurred during ceeded in wind-induced vortices some- periods of light winds, not exceeding where between Beaufort 2 and 3." They Beaufort force 2. On one occasion a noticed subsurface depletion of phos- patch disappeared when the wind rose phorus and high surface concentration from force 2 to 3. The blooms were during the 1953 Delaware Bay bloom.

58 which they state could be easily ac- p. 23-26. U.S. Comm.Fish Fish., Rep. counted for by current vortices extend- Comm., pt. 19, for year ending June 30,

ing down only 1 or 2 m. 1893.

". . , Before the influence of trace metals, vitamins, or other organic ma- RAY, SAMMY M., and WILLIAM B. WILSON. terials can be determined, improved 1957. Effects of unialgal and bacteria-free field methods for their quantitative es- cultures of Gymnodinium brevis on timation are needed. This remains one fish, and notes on related studies with of the most serious deficiencies in our bacteria. U.S. Fish Wildl. Serv., Fish. knowledge of the causative factors of Bull. 57:iii + p. 469-496; also U.S. Fish dinoflagellate bloonns." [p. 59.] Wildl. Serv., Spec. Sci. Rep. Fish. 211, iv +50 p. PORTER, JOSEPH Y. 1882. On the destructionoffishby poisonous Extensive experiments were per- water in the Gulf of Mexico. Proc, U.S. formed to prove that Gymnodinium breve Nat. Mus. 4:121-123. produces a toxin that kills fish. Ex- per indents showed that cultures of G. This report contains chiefly conjec- breve killed fish, while cultures of G. ture, attributing the fish kills to swamp splendens and of Prorocentrum sp. did water poisoned by dogwood, or to sub- not. marine springs. Fishermen catching Experiments with Flavobacteriuni fish north of Pine Island, Charlotte piscicida failed to show any toxic effect. Harbor, were losing fish in their live Lasker (Bein, 1954) had isolated this wells on the way to Havana. Concern- red-pigmented bacterium; Bein's ex- ing the submarine spring it is periments indicated 24-hour cultures of marine fish. stated, ". . .One proof of its volcanic killed several species orgin is that the water so polluted is of Experiments showed that water already a 'red brick color,' at a distance of nnade toxic does not lose its toxicity ." of less than a mile from the shore. . . for some time after the removal G. [p. 123.] breve . Likewise, the occurrence of any condition unfavorable to the organism POWERS, EDWIN B. may cause disintegration of the cells so 1938. Factors involved in the sudden mor- that the water may be toxic even though tality of fishes. Trans. Amer. Fish. it contains only fragmentary remains of Soc. 67:271-281. G. breve cells that are not readily identifiable. Filtrates of G. breve passed through PRATJE, A. a millipore filter under suction proved Noctiluca miliaris Suriray, Beitrage 1921. more toxic than filtrates passed by zur Physiologie, und Morphologie, gravity through filter paper. The authors Cytologie, I: und Morphologie suggested this difference may be due to Physiologie (Beobachtungen an der more organisms being retained intact lebenden Zelle). Arch. Protistenk. 42: by the filter paper. 1-98. [Non vidi.j They found no indication that fish kills by G. breve result from depletion PROVASOLI, LUIGI, of oxygen by the great masses of the 1958. Nutrition and ecology of protozoa and organism. algae. Annu. Rev. Microbiol. 12:279-308. The degree of toxicity of G. breve cultures was influenced, aside from Gives summary table of vitamin re- concentration, by such factors as the quirennents of numerous algae from growth phase of the culture, the pH of many authors; 165 references. the culture during the growth and the test periods, temperature and salinity of the test culture, the size and number A. McLAUGHLIN. PROVASOLI, L., and J. J. of the test fish, the volume and degree heterotrophy of some photo- 1963. Limited of aeration of the test culture, and In Carl H. synthetic dinoflagellates. bacterial growth. Oppenheinaer (editor). Symposium on "Bacteria-free cultures of G. brevis 105-113. marine microbiology, ch. 10, p. with concentrations varying from 2.3 Charles Springfield, 111. C. Thomas, to 4.8 million organisms per liter were toxic to two species of test fish. Five RATHBUN, RICHARD. species of fish were killed when sub- 1895. Mortality of oysters in Galveston jected to unialgal G. brevis cultures Bay. In Report upon the inquiry respect- containing 0.6 to 2.1 million organisms ing food-fishes and the fishing- grounds, per liter. . . ." [FB., p. 495.]

59 RICE, THEODORE R. in this field of research were invited 1954. Biotic influences affecting population by the Director of the Bureau of Com- growth of plamktonic algae. Fish Wildl. mercial Fisheries to a symposium on Serv., Fish. Bull. 54:ii+p. 227-245. red tide held at Galveston, Tex. It was the concensus of this group, after ma- "It is concluded that antagonistic sub- ture consideration of the copper sulfate stctnces arising fronn the metabolism experiment, that the combination of of phytoplankton are important, at least excessive cost, short duration of con- in fresh-water ponds, in influencing the trol, and possibility of harnn to other seasonal fluctuations in total phyto- marine life renders application of the plankton numbers and in the numbers method inadvisable. of each species, as well as in causing RYTHER, JOHN H. a definite succession of species." [p. 1955. Ecology of autotrophic marine dino- 244.] flagellates with reference to red water conditions. In Frank H. Johnson ROBINSON, REX J., and THOMAS G. THOMP- (editor). The SON. luminescence ofbiological systenns, p. 387-414. Amer. Ass. 1948. The determination of phosphates in Advance. Sci., Wash. D.C. sea water. J. Mar. Res. 7(1):33-41. "The dense populations of dinoflagel- The authors stated, "If the samples lates which create 'red water' conditions of sea water contain much plankton are known only in the tropics or in this should be rennoved by filtration temperate water during the warnner or centrifugalization. However, it is (and usually the warmest) time of the

only in extreme cases when such a year. . . . procedure is necessary." [p. 38.] Not "Such fragmentary physiological evi- discussed are errors in phosphorus dence as is available concerning the determinations from particulate phos- temperature relations of dinoflagellates phorus in unfiltered samples. appears to support the view that they are predominantly a warm- water group. ROTBERG, M. Barker (1935), who is one of the pioneers 1958. The thiannine requirements of in developing successful culture methods Prymnesium parviim (Chrysomonadina). for dinoflagellates, observed optimal Bull. Res. Counc. Israel 7B:208-210. temperatures for the growth of some 14 species between 18° and 25° C. ROUNSEFELL, GEORGE A. Braarud and Pappas (1951) noted a 1958. Large-scale experimental control of temperature optimum for Peridinium

red tide. In Annual report of the Gulf triquetrum at 18° C, while Nordli ( 1 953) Fishery Investigations for the year end- found that Ceratium fusus and C. furca ing June 30, 1958, U.S. Fish Wildl. Serv., grew most rapidly at temperatures of p. 76-77. 15° and 20 C respectively. Provasoli (personal communication) finds tem- Information herein published by peratures of 20-25° C most suitable for ." Rounsefell and Evans (1958). growing Gyrodinium californicum . . . [p. 389-390.] ROUNSEFELL, GEORGE A., and JOHN E. The author gave the following table EVANS. to show preferred salinities: 1958. Large-scale experinnental test of cop- per sulfate as a control for the Florida Optimum and Maximum Range of Salinity for Groirth of Some Neritic Dinoflagellates red tide. U.S. Fish Wildl. Serv., Spec. Sci. Rep. Fish. 270, vi + 57 p.

Gives results of the first large-scale attempt to control overblooming of the

red-tide organism, Gymnodinium breve .

". . . About 16 square miles stretching along 32 miles of shoreline from Anclote Key to Pass-a-Grille Beach, off St. Petersburg, Florida, were dusted with copper sulfate (CUSO4.5 H2O) at about 20 pounds to the acre by crop -dusting

planes. . . ." [p. iii,] In March 1958, after the completion of the manuscript of this report, a selected panel of 14 biologists versed an extremely wide range of salinities. stimulated the growth of Peridinium This high degree of adaptability is a triquetrum ." [p. 393.] definite advantage to life in the variable The author suggested that growth environment of coastal and estuarine of dinoflagellates may be dependent waters, and it is perhaps one of the upon, or at least benefited by, the means by which the neritic dinoflagel- previous flowering of diatoms. This lates are able to compete successfully he suggested (citing Pearsall, 1932, with other organisms, such as diatoms, and Hutchinson, 1 944) as possibly caused which in general have a nnore narrow by the reduction of the concentrations range of salinity tolerance." [p. 391.] of one or more of the nutrients or The author, citing Fritsch (1935), trace metals by diatoms to a level mentioned that the naked dinoflagellates favorable for dinoflagellates. On the appear to be most abundant in the open other hand, Lucas (1947, 1949) pro- ocean plankton, while the armored forms posed that the production of external are more typical of coastal and estuarine metabolites, or "ectocrines" by one regions. group of plankton organisms may bene- "The dinoflagellates have often been fit the succeeding population, but in- credited with the ability to utilize and hibit competing organisms. flourish in extremely low concentra- "The dinoflagellates . . . possessing tions of nitrogen and phosphorus (Gran, the advantage of motility, are able to 1926-27; Gilson, 1937). This concept maintain themselves in water of low has stemmed largely from observations density with comparative ease and are that dinoflagellate maxima, intemperate relatively independent of vertical nnix- waters, follow the decline of the spring ing. ... [p. 395.] diatom flowerings and relatively large ". . . motility provides the dinoflagel- populations often persist throughout the lates with an advantage in waters of summer months when the supplies of low nutrient content. The nonmotile these nutrients are almost undetect- diatoms are dependent upon the dis- able. Gran (1926-27) has proposed that solved nutrients contained in the water the dinoflagellates require less nutrition which immediately surrounds them and for growth than the diatoms on account through which they sink (see Munk and of their relatively low rate of nnetab- Riley, 1952). In contrast, the dino- olism. [p. 391-392.] flagellates, thoughnot strong swimmers, can move about for considerable dis- tances and localize in the most ad- vantageous depth for .

". . . dinoflagellates as a group show If the nutrient level of the water is a continuous variation in their modes low, they may, by their vertical migra- of nutrition from autotrophic to holozoic, tions, utilize all the nutrients available while mamy species are facultative, ob- within the entire photic zone. Accord- taining their food by either or both ing to Peters (1929) Ceratium can move methods (Kofoid and Swezy, 1921), . . . through 5 to 10 meters in 12 hours or [p. 392.] less." [p. 397.] The report contains an extensive table of 21 red-water occurrences in various parts of the world, including "In inshore waters dinoflagellates some that caused nnass mortalities. are often abundant, and in many parts The author stated, ". . . one factor which of the world their populations may de- is alnnost universal in red water out- velop bloom proportions. Here the close breaks, the occurrence of a high water association of the plankton with land temperature. In temperate or boreal masses and the contribution of runoff regions of the ocean, red water appears water to their environment provide to be restricted to the summer months, what may be considered as a natural and the notation is frequently made 'soil extract,' and, in the vicinity of that it is preceded by periods of un- heavily populated areas, frequently a usually hot, calm weather. Along the source of organic pollutants. As men- Indian coast, red water occurs during tioned earlier, Braarud (1945) observed the clear, hot periods between the a heavy growth of dinoflagellates in southwest and the northeast monsoons the highly polluted regions of the (Hornell and Nayudu, 1923; Menon, 1945; Oslofjord. Braarud and Pappas (1951) Bhimachar and George, 1950). Off the later found that the addition of small Peruvian and Southwest African coasts amounts of raw sewage to the medium it appears during the southern summer

61 when the upwelling of cold water is two such adjoining cells. Under these at a minimum and water temperatures conditions parallel streaks of floating are the highest of the year (Brongersma- matter are produced (Fig. 5). (For a Sanders, 1948). ... [p. 401.] more detailed description of this proc- ess, see Langmuir, 1938; Stommel, 1949). [p. 406-407.]

". . . there may be insufficient nutri- ents normally present in sea water, and no known mechanism for concen- "Thus there is no necessity to postu- trating them to a sufficiently high level, late obscure factors which would account to support the development of a typical for a prodigious growth of dinoflagellates red tide. The remaining possibility, to explain red water. It is necessary that the organisms themselves become only to have conditions favoring the concentrated after growth, will be dis- growth and dominance of a moderately cussed below." [p. 403.] large population of a given species, The author here discussed at length, and the proper hydrographic and mete- with much controversial documentation, orological conditions to permit the ac- the vertical migration of dinoflagellates cumulation of organisms at the surface and how light and buoyancy can affect and to effect their further concentrations it. His evidence for surface concentra- in localized areas." [p. 409.] tion is sketchy (for only two species in 6 m. of water). He said the accumu- RYTHER, J. H., and R. R. L. GUILLARD. lation of buoyant organisms at the 1962. Studies of nnarine planktonic diatoms. surface would be enhanced in the ab- II. Use of Cyclotella nana Hustedt for sence of vertical mixing of the water, assays of vitamin B12 in . and that most occurrences of red tide Canad. J. Microbiol. 8(4): 437-445. throughout the world were accompanied or preceded by periods of calm weather SALSMAN, G. G., and W. H. TOLBERT. and smooth seas. He added that such 1963. Surface currents in the northeastern conditions, together with the high tem- Gulf of Mexico. Navy Mine Defense peratures which usually accompany Lab., Panama City, Fla., Rep. 209,43p. them, may cause thermal stratification, giving additional resistance to vertical SATER, EDNA N. mixing. 1954. Florida's red tide problem. Fish Ryther suggested three means by Wildl. Serv., Fish. Leafl. 420, 11 p. which organisms accumulated at the surface of the water nnay be further This leaflet reviews generallythe red concentrated: tides from 1946-54 and research by "(1) Prevailing onshore winds: Sur- Federal, State, and private agencies. face water driven shoreward by pre- The theory is discounted that alleged vailing onshore winds establishes a dumping of war munitions brought about circular pattern, sinking at the waters the kill of fish in 1946 and 1947. edge and returning seaward at lower "Red tides occur on the west coast depths. Buoyant organisms will ac- of Florida only when certain peculiar cumulate in windrows along shore or conditions prevail: after a period of at the region of descent (Fig. 3). abnornnally heavy rainfall followed by a "(2) Where brackish coastal water, shift from offshore to onshore winds. particularly in the vicinity of river These circumstances lead to the accu- mouths, meets open ocean water, there mulation of a mass of water of ab- is a mixing and sinking of the two water normally low salinity, which is kept masses along a line of convergence. from dispersing seaward by the winds Both types of water flow toward this blowing toward shore. In this water line, and buoyant organisms will ac- mass, the organism G. brevis explodes cumulate at or near the convergence into what biologists call a bloom {an line, producing streaks of floating ma. extraordinary increase innumbers), and terial (Fig. 4). becomes poisonous to fish life. As the "(3) Convection cells: Wind-driven fish die, their decaying bodies release vertical convection cells may be estab- nutrients which nourish the bloom and lished which rotate alternately clock- intensify it. [p. 6.] wise and counterclockwise with their vertical axes perpendicular to the di- rection of the prevailing wind. Floating objects will accumulate in the region "The theory is now held that the red- between the descending connponents of tide organisms are able to multiply

62 rapidly because nutrients are received of the Gulf States Marine Fisheries ." through drainage from the land. . . Commission. The work of the three ." [p. 7.] agencies is thus coordinated. . . In 1948, a small Federal laboratory- [p. 10-11.] was established at Sarasota, where laboratory culture experinnents and SCHILLER, JOS. hydrographic sampling at sea were 1933. Dinoflagellatae. In Kryptogamen- undertaken. While the laboratory was Flora (L. Rabenhorst). Band 10, Abt. at Sarasota, from 1948 to 1952, how- 3, Teil 1. Akad. Verlags., Leipzig, ever, not one specimen of G. breve 617 p. was taken in samples. The laboratory moved back to Galveston in June 1952, SCHILLER, JOS. and the red tide broke out again in 1937. Dinoflagellatae. In Kryptogamen- November. Flora (L. Rabenhorst). Band 10, Abt. Bureau scientists were able to keep 3, Teil 2. Akad. Verlags., Leipzig, G. brevis cultures alive after the 1952 589 p. outbreak. The Alaska collected a great deal of hydrographic data in the red- SEYDEL, ElvUL. tide area, however, and had some 1913. Fischsterben durch Wasserbliite. success at killing red-tide organisms Mitt. Fisch.-Ver., Prov. Brandenburg, by discharging a concentrated solution N.F. 5(9):87-91. of copper sulfate from its ballast tanks into the ciffected areas. SHERWOOD, GEORGE H., and VINAL N. Another outbreak followed in Sep- EDWARDS. tember 1953. "During this outbreak, 1902. Red tide in Narragansett Bay, R.I. copper-sulfate crystals were placed In Biological notes no. 2 (a contribution in sacks and towed behind small boats, from the U.S. Fish Commission Biologi- off Anna Marie Key, at the mouth of cal Laboratory, Woods Hole, Mass.), Tampa Bay. Again this experiment was p. 30. U.S. Fish Comm., Bull. 21.

successful in killing the organisms. . . . [p. 8.] SHILO, MOSHE, S, SARIG, MIRIAM SHILO, and H. ZEEV. 1954. Control of Prymnesium parvum in fish ponds with the aid of copper sul- "From 1948 the Federal Government fate. Bamidgeh, Bull. Fish Cult. Israel spent approximately $50,000 each year 6(3): 99-102. on red-tide research, until 1953 when the appropriation was reduced to SHILO [SHELUBSKY], M., and M. ASCHNER. $35,000. In the fiscal year 1955, $70,000 1953. Factors governing the toxicity of cul- has been earmarked for the Fish and tures containing the phytoflagellate Wildlife Service's use. In January 1954, Prymnesium parvum Carter. J. Gen. the Service reestablished a field station Microbiol. 8(3):333-343. [Reference in Florida, this time at Fort Myers .... from Ray and Wilson, 1957.]

Concerning this paper Ray and Wilson (1957, p. 488) stated: "Another role "In May, a new 43-foot vessel, since for which bacteria must be considered named the Kingfish , was delivered at is that of a detoxicating agent. Shilo Fort Myers for use in studying the and Aschner (1953) found that bacteria ocean conditions that cause red decreased the toxicity of cultures of tides. ... [p, 9.] Prymnesium parvum, a marine and brackish water chrysomonad that is toxic to fish. Similarly, bacterial ac- tivity may influence the toxicity of

". . . the Gulf States Marine Fish- C. brevis in the laboratory and in eries Commission, established by a nature." See also McLaughlin (1956). compact among the five Gulf States, has appointed a special red-tide com- SLOBODKIN, L. BASIL. mittee, of which Dr. F. G. Walton 1953. A possible initial condition for red Snnith, Director of the University of tides on the coast of Florida. J. Mar. Miami Marine Laboratory, is chairman. Res. 12(1):148-155. On the committee are representatives of the University of Florida at Gains- A mininnum critical size is assumed ville and of the Fish and Wildlife Serv- for a water mass physiologically suit- ice, by law the primary research agency able for the growth of a phytoplainkton

63 population, which will permit an in- To these reviewers this is not con- crease in the concentration of orga- firmed by the data of the Alaska (Collier, nisms. It is also stated, however, that 1958a). The station referred to was this relation is not of particular signifi- located at lat. 28O30' N. and long. 84°32' cance if the phytoplankton species in W. (station 36). We have prepared a question is tolerant of a wide range of table showing pertinent data for this environmental conditions. The basis and the "surrounding" stations, in- used to assume intolerance is two per- cluding the three (Nos. 35, 38, and 40) sonal communications, one from Luigi to which Slobodkin referred. The nearest Provasoli stating that G. breve is not station was about 40 miles distant. It generally amenable to culture tech- may be noted further that two of the niques, and the other from Herbert three salinities to which he referred Graham stating that G^. breve is not were not at the surface but at 10 m. normally found in the marine phyto- Furthermore, at the station in question plankton of the region. We do not accept (No. 36), the salinity chcinged (?) from these two premises. Furthermore, the 33.42 to 33,48 p.p.t. in 42 minutes. assumption that red tide is found in very discrete water masses is at vari- ance with the statement of most field 1952 observers that red tide water occurs in streaks, not in regular-shaped masses. As confirmation of his theory, Slo- bodkin mentioned red discoloration of the sea observed at lat. 28°30' N. and long. 84°30' W. from the vessel Alaska on June 2, 1952. He stated, ". . . This discoloration was caused by a dense population of a red ciliate, probably Mesodinium pulex Noland or a closely related form (Kudo, 1947). The surface salinity at a hydrographic station in the discolored water (33. 4%o) was lower than that at surrounding stations (34.99, 35.47, 36.17). This would seem to con- firm the theory in so far as protista in general are concerned, since the dis- colored water was approximately 90 miles from the nearest coast." [p. 149, 151.] TABLE 1(a) Continued — Location of Rainfall (inches), Month;' Year Date of Initial Report Red Tide Weather Station

1908 Reported by Taylor (1917) as killing sponges, without any mention of dead fish. This is probably an outbreak of sponge blight rather than red tide (Smith, l':*'^!) 1916 October 3 (Taylor, 1917) Boca Grande to 12.56, July; 8.22, August; San Carlos Bay 5.34, September; Fort %-ers 1946* November 20 (Gunter, et al., Naples 9.23, September; 2.03, 1948) October; 9.69, November; Naples In October 1946, Fort Meyers, Punta Gorda and the Everglades, Florida were flooded by unusual tides in the wake of a hurricane on October 7-8. Charlotte Harbor and part of the Everglades can be considered as having been thoroughly mixed with sea water, which drained into the Gulf of Mexico in considerable volume. November was warmer by 6.1° than any previous November on record (71.2° mean for Florida). October was 1.1° above normal and warmer than any October in five years for the state as a whole.

1947* March (Gunter, et al., 1948) Florida Bay 10.37, March; Captiva 8.94, March; Fort Meyers Greatest March rainfall on record for state with ex- ception of 1930 1947*t June 20 (Gunter, et al., 1948) Bonita Beach 6.47, May; 12.84, June; (South of Fort Fort %-ers (2.69 inches %ers) fell after June 20) 3.34, May; 13.33, June; Punta Gorda (.04 inches fell sifter June 20) 1947*t July 18 (Gunter, et al., 1948) Venice to 13.06, June; 11.44, July; Sarasota Sarasota (7.64 inches after July 18) 18.25, July; Punta Gorda (12.72 after July 18) Floods in Okeechobee region diiring June and July 1952 October 25 Boca Grande and 12.35, September; 8.34, Southwest of October; Fort %ers (none Sanibel after October 25)

*A11 outbreaks marked with an asterisk are considered by Gunter, et al. (1948) as representing a single occurrence. Since discoloration and fish mortality disappeared between each of the listed dates, they are here considered as separate. tThe interval with no fish kills separating these two outbreaks is of the order of two weeks and may or may not indicate separate dinoflagellate populations."

Slobodkin then presented a table for comparison with table I (a) above.

"TABLE 1(b). --Mean Rainfall (inches) by Months at Five Florida Weather Stations SMITH, F. G. WALTON. SMITH, F. G. WALTON. 1949. Probable fundamental causes of red 1957. Mystery of the red tide. Smithsonian tide off the west coast of Florida. Inst., Annu. Rep. for 1957:371-380. Quart. J. Fla. Acad. Sci. 11(1): 1-6. This is a semipopular article without The theory is advanced that the red- documentation, apparently largely a re- tide blooms must depend upon unusual print from Sea Frontiers 3(1):21-31. amounts of phosphorus. The origin of Some of the illustrations appeared in such phosphorus could be deposits in Galtsoff (1949). river drainages or offshore deposits On page 373 it mentions "as many but the manner of dispersal remains as 60,000,000 individual cells to the a nnystery deserving investigation, pint of water" but we are sure this is merely an error in transposing a SMITH, F. G. WALTON. liter into an English equivalent.

1954. Emergency report on the Florida red ". . . Since 1947 red tide seemed to tide, January 1954. Univ. Miami, Mar. have disappeared and there was no way Lab., Rep. to Fla. State Bd. Conserv., of telling whether it might return in Tech. Rep. 54-2, 4 p. 1 year or 10 years. ... [p. 376.]

This report sumnnarizes the status of red-tide research by the Marine Labo- ratory as of January 1954. It names "These expectations were partly Gymnodinium brevis as the causative realized in 1952 when a fresh but minor organism of the Florida red tide, aind outbreak occurred. About the middle of says that red tide occurs sporadically; September 1953 further red tide was damage to commercial fisheries in a reported and this continued at inter- bad red-tide year exceeds $1 million; vals throughout the winter and in the Gymnodinium brevis releases fish- spring and summer of 1954. The new killing into the water; phos- alarms brought special funds to aid phorus compounds are necessary for research at Miami and increased fed- plamkton growth; other factors besides eral activity. The State of Florida phosphorus are necessary for the red made a wise move by setting up a Red- tide; attempts to establish correlations Tide Committee in order to coordinate

with meteorological phenomena were research activities. . . . unsuccessful; and laboratory culturing "Materials are needed for the growth of Gymnodinium provided information of 'Jim brevis' and the suspicion that on special nutrient requirements. the brackish bay waters contained sonne The author believed that destruction essential part of these materials re- of the bloom serves little purpose sind ceived new attention as the result of that the only solution is the prevention work carried out by the Haskins Lab- of bloon-iing. oratory in New York. For the first A 2-year program at a cost of $72,000 time the red-tide type of organism was per year is outlined to ensure success- kept alive in the laboratory in a pure ful prediction, prevention, and control culture, uncontaminated by bacteria or ." of red-tide blooms. This program en- other organisms, . . [p. 377.] (Re- tails: viewers' note: These were dinoflagel- lates other than G. breve, which was "(a) Full-time year-round oceano- first cultured in a pure state at Galves- graphic investigation of the chemical ton by the Fish and Wildlife Service.)

cuid physical changes of the West Coast ". . . after taking meteorological fig- waters. ures for 26 past years and performing "(b) Multiple statistical correlation numerous calculations with different of outbreaks with meteorological and combinations of the data, a formula other ambient phenomena. emerged which worked. The weather "(c) Tropistic and nutritional labora- information for any year was placed tory studies of the causative orga- into the formula. When the numerical nism." [p. 4.] result fell within a certain narrow The author stated that the greatest range, then a red-tide outbreak hap- success will be obtained by providing pened during the next 12 months. If adequate funds which "should be con- outside the range, there was no red centrated in one locality or organiza- tide. tion, equipped with proper laboratory "... In November 1955 the State facilities and ocecinographic vessels, Board of Conservation in Florida was and with personnel experienced in the notified that there was little likelihood Red Tide problem." [p. 4.] of major red-tide outbreaks in the year

66 -

1956. It turned out that there was none. Toxin was prepared from cultures of A similar prediction was made for 1.5 nnillion living cells of G. breve p.p.t., 7 1957. . . ." [p. 379.] (Reviewers' note: (at a pH of 8.15, salinity 33 A heavy outbreak occurred in 1957.) weeks old) frozen overnight in flasks, thawed at room temperature (25+ 2°C.) SOLI, GIORGIO. and filtered through an AA millipore 1964. A system for isolating phytoplankton filter. Dilutions were made with sea- organisms in unialgal and bacteria-free water medium of 33 p.p.t., pH of 7.9. culture. Limnol. Oceanogr. 9(2): Preliminary experiments with guppies 265-267. and mullet indicated that living cultures of G. breve were less toxic than killed SOMMER, HERMANN, and FRANCES N. ones. The toxin preparation was tested C LARK. on mullet at several dilutions and the 1946. Effect of red water on marine life results were shown in a curve. If we in Santa Monica Bay, California. Calif. interpret his curve in numbers of G. Fish Game 32(2): 100- 101. breve per liter needed (when killed) to produce the effect, the results are During June 1945, red water occurred approximately: from San Luis Obispo to Harbor. On June 19 dead fish and shell No. of cells Average time of death fish--chiefly spiny , Panulirus inte r ruptus - -we re observed in Santa 1,500,000 2 to 4 minutes Monica Bay. Also affected were spider 450,000 under 30 minutes crabs and many fishes, including halibut, 300,000 over 50 minutes stingrays, and sharks. Ceratium was 150,000 4 to 8 hours, sometimes suspected as the cause but proof was failed to kill not definite. Residues (cell debris) left onAAmil- SOMMER, HERMANN, W. F. WHEDON, C. A. lipore filters contained little or no toxic- KOFOID, and R. STOHLER. ity. Heat had an effect on toxicity of the 1937. Relation of paralytic shell-fishpoison cell-free toxin preparation. Whenheated to certain plankton organisms of the and cooled immediately in an ice bath, genus Gonyaulax. Arch. Pathol. 24(5): little toxicity was lost from heating to 537-559. 80° C, but most of the toxicity was lost from heating to 100° C. SPECHT, R. C. Heating over longer periods destroyed 1950. Phosphate waste studies. Fla. Eng. toxicity at lower temperatures. Thus, Ind. Exp. Sta., Bull. 32, 27 p. most of the toxicity was lost by heating to 45° C. for 4 hours, and no toxicity Figures are given showing the phos- was detected after heating to 100° C. phate content of the Peace and Alafia for 5 minutes. Rivers to be very high. Toxicity of living unialgal cultures was increased by pH of 8.1 (5-weeks- SPENCER, C. P. old culture, salinity 30 p.p.t., 1.2million 1952. On the use of antibiotics for isolating per liter) being changed by adding 5N bacteria-free cultures of marine phyto- HCl or 5N NaOHtothemedia. AtpH 3.3, plankton organisms. J. Mar. Biol. Ass. 5.4, and 9.5, G. breve were destroyed U.K. 31(1):97. and toxicity was, thus, increased. in pH of cell -free toxin had STARR, Changes THEODORE J. little or no effect on toxicity. Adding vitamin iri 1956. Relative amounts of B12 sufficient heavy metals to kill G. breve estuarine detritus from oceanic and increased the toxicity of unialgal cul- Island, environments near Sapelo tures. Georgia. Ecology 37(4):658-664.

Mentions found in bottom mud. B12 STOHLER, R. i960. Fluctuations in moUusk populations STARR, THEODORE J. after a red tide in the Estero de Punta 1958. Notes on a toxin from Gymnodinium Banda, Lower California, Mexico. breve . Tex. Rep. Biol. Med. 16(4): Veliger 3(l):23-28. [Cited from Biol. 500-507. Abstr.] Starr described procedures for the bioassay of the toxin in unialgal cul- A collection of mollusks made during tures of G. breve and some properties the peak of a red tide in 1958 was com- of crude toxin preparations. pared with collections made in July

67 1959 and January 1960. It was con- The only other active principle came cluded that neither redtide nor seasonal from cells of another dinoflagellate,

variations could be held responsible P ro roc ent rum micans , also grown in for the fluctuations observed in sizes 4 percent soil extract. and occurrence of specimens. SWEENEY, BEATRICE M. 1954. Gymnodinium splendens STOMMEL, HENRY. , a marine dinoflagellate requiring vitamin 1949. Trajectories of small bodies sinking Bi2' Amer. J. Bot. 41{10):821 -824. slowly through convection cells. J. Mar. Res. 8(l):24-29. [Cited by Ryther, SWEENEY, BEATRICE M., and J. WOODLAND 1955.] HASTINGS. 1958. Rhythmic cell division in populations

Explamation of the accumulation of of Gonyaulax polyedra . J. Protozool. floating objects in windrows parallel 5(3):217-224. to the wind direction by wind-driven convection cells. "In cultures of the marine dinoflag- ellate Gonyaulax polyedra grown with STREETS, THOS. H. alternating light and dark periods of 1878. The discolored waters of the Gulf of 12 hours each, at least 85% of all cell California. Amer. Natur. 12(l):85-90. divisions which occur in a day take place during a 5-hour period spanning the end STRODTMANN, S. of the dark period and the beginning of 1898. Ueber die vermeinte Schadlichkeit the light period. A very distinct maxi- der Wasserbliite. Forschungsber. Biol. nnum in the number of recently divided Sta. Plon. 6(9):206-212. cells occurs at about the time the light period begins. SWEENEY, BEATRICE MARCY. "This diurnal periodicity in cell divi- 1951. Culture of the dinoflagellate Gym- sion is lost after 4 to 6 days in con- nodinium with soil extract. Amer. J, tinuous bright light, but will persist for Bot. 38(9):669-677. at least 14 days in continuous dim light. The period of approximately 24 hours Soil extracts have been widely suc- under constant conditions is only slightly cessful in promoting the culture of altered by varying light intensity and organisms difficult to grow in inorganic tennperature, both of which factors are media. The active principle in soil known to affect markedly the generation extract according to Pringsheim (E. G. time. The tinne at which cell division Pringsheim, 1936, Das Ratsel der occurs is therefore postulated to be Erdabkochung. Beih. Bot. Centralblatt controlled by some sort of rhythmic A55: 101-121) in experiments using or clock mechanism.

Chlorogonium , Polytoma , Polytomella , "Similarities between the cell divi- £ind Chilomonas is am acid- and alkaline- sion rhythm and the endogenous rhythm stable organic substance, insoluble in of luminescence in this organism are alcohol and ether, adsorbed by charcoal, discussed. The occurrence of a glow, and destroyed by hydrogen peroxide. or steady light emission of low intensity, Gymnodinium splendens was cultured at about the time of cell division, is in several media, usually in inorganic described, and it is suggested that this media with organic additives. No in- glow may result from cellular changes organic additives aided growth. Soil accompanying certain stages of cell extract promoted growth. The soil ex- division." [p. 217.] tract was made from redistilled water and garden soil, which was autoclaved, TABB, DURBIN C., and DAVID L. DUBROW. decanted after standing overnight, and 1962. Hydrographic data, Supplement I, reautoclaved. Although presumably bac- from the inshore bays amd estuaries teria-free, the soil extract was of little of , Florida, or no avail when first prepared; it 1959-1962. Univ. Miami, Mar. Lab., reached a peak of activity (in promoting Rep. to Fla. State Bd. Conserv. 62-9, growth) in 4-6 weeks and declined in 22 p. activity after about 80 days. Activity Includes tennperatures, salinity, oxy- of soil extract was not retained by re- gen, and pH. autoclaving or by refrigeration at 4° C. Deep freezing at -25° C. preserved ac- TABB, DURBIN, DAVID DUBROW, and RAY- tivity for 6 months. Trials of additions MOND MANNING. of several different amino acids, yeast 1959. Hydrographic data from the inshore extract, oak leaf mold extracts, etc., bays and estuaries of Everglades Na- were unsuccessful. tional Park, Florida, 1957-1959. Univ.

68 Miami, Mar. Lab., Rep. to Fla. State other water birds behaved normally. Bd. Conserv. 59-5, 26 p. Buzzards were common but neglected the fish entirely.

". . of the mortality, TAYLOR, HARDEN F. . at the height 1917a. A mortality of fishes on the west on the Gulf coast, the water was of an

coast of Florida. Science 45{ 1 163):367- amber color (by transmitted light). This 368. [Cited from Gunter, 1947.] colored water was described as being not uniformly distributed, but occurring in TAYLOR, HARDEN F. streaks, and it was in these streaks 1917b. Mortality of fishes on the west that the fish are said to have per- coast of Florida. Rep. U.S. Comm. ished. ..." [p. 11.] Fish, for 1917, append. 3, 24 p. (Doc. TORREY, HARRY BEAL. 848.) 1902. An unusual occurrence of Dinoflagel- lata on the California Coast. Amer. ". . The reports and references . Natur. 36(423): 187-192. are too fragmentary to give an accurate

record . . . but collectively they clearly Bloom was first noted on July 7, 1901, indicate that all the keys from Key as a red streak off the mouth of San West nearly as far north as Cedar Pedro Harbor. He identified it as Gon- Keys have been visited by this plague, yaulcLX and mentioned am unusual display and that it occurred in the years 1844, of phosphorescence. The "muddy ver- 1854, 1878, 1880, 1882, 1883, 1908, and nnilion" streak reached the shore on the finally in 1916. l6th. On the 20th the beach (about 400 "In October and November, 1916, yards long) was lined with dead the mortality recurred in severe form, animals - -several hundred holothurians,

the first visitation since 1908. . . . stingrays, guitarfishes, sharks, red "Fishes of a great number of species perch, smelts, and several octopi. were noted dead and dying; the air was The "red water" occurred for 200 charged with a suffocating gas, miles, from Santa Barbara to San Diego, and extended several miles to sea. It which . . . irritated the air passages, producing the symptoms of colds. This was still present around Sam Pedro on gas, while exceedingly irritating, had September 1.

". . in great no odor. . . . The abnormal conditions . Noctiluca appeared seemed to be moving southward, oc- numbers toward the end of July, cind ." curring at Boca Grande on October 3 devoured Gonyaulax with avidity. . and 18, at about the He mentions nine additional species of middle of October, at dinoflagellates present in lesser abun- about October 20, at San Carlos Pass dance. The red water was not present about November 1, and dead fish were at Santa Catalina Island, 20 miles off- first seen at Big Marco Pass onNovem- shore. ber 5. Captiva Pass is 7.5 statute miles "The boundaries of the red streaks south of Boca Grande Pass; the others were quite sharply marked, although the are, respectively, 16.5, 27.75, and 67.75 water between streaks often contained ." miles to the south of Boca Greinde Gonyaulax. . [p. 191.]

Pass. . . . dead fish were seen as far U.S. FISH and WILDLIFE SERVICE. south as Cape Romano. . . . The death 1958. Red tide symposium, Galveston, of two persons in Fort Myers, Fla., in Texas, March 5-7, 1958. 72 p. November, was attributed to the eating of some of these dead fish." [p. 5-6.] U.S. FISH and WILDLIFE SERVICE. He quoted a letter from the deputy 1961. The Florida redtide. U.S. Fish Wildl. collector of customs at Boca Grcinde Serv., Fish. Leafl. 506, 8 p. [Revision stating that the dead fish started to of FL 420.] come ashore on October 3. In a few During the 1 946-47 outbreak: --"Prac- days the plague abated, but another tically all species of fish, including violent outbreak occurred on the 18th, such large forms as tarpon andjewfish, killing larger fish cind many of the bay were included in the victims of the red fishes. The gas was worse, and many water. Most oysters in affected areas people asked for medical assistance. died. Horseshoe crabs died by the listed 63 species of fish Taylor thousands, but true crabs apparently killed and sea urchins (Arbacia), horse- were unharmed. Sponges showed no shoe crabs, and sponges. He said bar- ill effects (the principal sponge beds nacles, oysters, and mussels were in near Tarpon Springs were outside the good condition, live conchs and hermit Red Tide area), 2.] crabs were observed repeatedly, por- [p. poises were plentiful, and pelicatns cind

69 "In 1953, Red Tide organisms at the dry spell, and quit in September 1947, Galveston Laboratory were first cul- during the heaviest rainfall, ." tivated artificially. . . [p. 6.] "6. The results of hydrographic studies carried out in previous years UNIVERSITY OF MIAMI, MARINE LABORA- had demonstrated the presence of Red TORY, Tide at the interface between Gulf smd 1954. Red tide studies, January to June Bay waters. Attempts were made to 1954. Prelim. Rep. to Fla. State Bd. develop further these relationships, but Conserv. 54-19, 117 p. no evidence was found to substantiate the idea of a more or less continuous This report covers many aspects of front between Gulf and Bay waters along red-tide research carried out by the which the Red Tide progresses. It now Marine Laboratory, but also sum- seems far more probable that a pe- nnarizes research by other agencies, riodic separation of masses of Bay without specifically mentioning under water susceptible to Red Tide takes whose auspices the work was accom- place at the mouth of the passes and plished. Despite an occasional refer- that these masses move with prevailing ence to an outside author, there is no currents, usually northward, slowly bibliography or list of references cited. losing their identity by mixing and Because of this lack of documentation diffusion with the Gulf Water." [p. 1-2.] and a list of 1 9 personnel (total of only We do not accept this latter theory 866 man days, including boat opera- (see Chew, 1953).

tions), the report is clearly intended "7. , . , initial fish kills are most

(as the title indicates) as merely a active and . . . G_. brevis blooms nnost progress report. It is stated that, "This often originate at the inner side of such is ." report a preliminary account of the passes as Boca Grande. . . [p. 2.] results. Because of the emergency na- "10. A study of commercial fish ture of the Red Tide problem, and, in landings fails to reveal any significant order that the data may become imme- decrease in the commercial catch in diately available to other workers in the relation to Red Tide. ... [p. 2.] field, it has been prepared for a re- "12. It is strongly recommended that stricted distribution without the careful advance planning be undertaken for the evaluation normally given to scientific control of Red Tide should outbreaks reports. For this reason it is in many occur in the fall of this or future years. ways incomplete and all discussions smd The most practical method of alleviating conclusions must be considered as ten- the damage is consideredto be by seining tative in nature, subject to re -evaluation of dead fish while concentrated in the as the data receives more careful passes and at sea, before drifting on scrutiny." [p. 3,] the beaches. The report starts with a summary: "It is considered that chemical con- "4. In the course of the work a sur- trol is more expensive and less likely vey was made of the scientific litera- to be successful than the control of dead ture concerning plankton bloonns fish by seining. Nevertheless, it is rec- throughout the world." [p. 1.] This onnmended that chemiical treatment be survey has not been published. applied at the passes at the earliest "5. . . . The initial series of fish possible moment of a Red Tide out- kills constituting a Red Tide cycle break in order to evaluate its useful- were found to occur most frequently ness." [p. 2.] Concerning treatment in in the month of October and within the passes see Ingle et al. (1959). dates of the new moon plus or minus "13. ... it is suggested that attention three days." [p. 1.] This conclusion be paid to the possibility of modifying is not accepted by us as having been the nutrient conditions of the Bay waters proved. in such a way as to prevent Red Tide "A cycle or series of outbreaks is conditions from developing. A large mostly likely to occur when the amnual scale fish culture program in the Bays, rainfall of the Peace River drainage with the addition possibly of supple- area is above the fifty year average. mentary nutrients other thain phosphorus A correlation also exists between out- might conceivably be brought about. breaks and a high maximum annual Under these conditions the nutrient discharge of the Peace River." [p. 1.] regime and the food chains might con- No actual statistical analysis is shown, ceivably be chcLnged so as to minimize and the best example shown of high the development of conditions suitable rainfall, in 1947, gives a false im- to Red Tide." [p. 2.] pression since the 1947 outbreak really On page 3 it is stated that "During started in November 1946, after a long the 1947 outbreak, scientists of the

70 Marine Laboratory of the University vert the suspected forms to the identi- of Miami established that the fish mor- fiable motile stage were unsuccessful. tality and the irritant gas associated with it were due to a plankton that even bloom. . . ." Woodcock (1948), from the "It was frequently noted Woods Hole Oceanographic Institution, when no other orgamism was present, a established this connection between red bloom of G. brevis contained large tide and the irritant gas carried as an numbers of the copepods Acartia ." aerosol. sp. . . [p. 74.] On page 7 it is stated, "The theory of submarine springs as a possible indirect UNIVERSITY OF MIAMI, MARINE LABORA- cause for the Red Tide has been tested; TORY. however, the results, so far, seem to 196 la. A report of data obtained in Florida show that this explanation is not cor- Straits and off the west coast of Florida, rect." There is no explanation of what January- June I960. Rep. to Off. Nav. "test" was applied. Res., Tech. Rep. 6I-I, 189 p. "The annual phosphate production A report containing raw hydrographic (land pebbles) statistics were studied. data obtained in an area north of Key No correlation was found between year West from Cape Sable to Dry Tortugas to year fluctuations in the phosphate (January-June, 1960). production and the outbreaks of Red Tide." No data are presented. UNIVERSITY OF MIAMI. MARINE LABORA- ". . of the method of least . By means TORY. squares it was found that the total phos- 1961b. A report of data obtained in Florida phorus content in the Peace River water Straits and off the west coast of Florida, is around 31 and in the Caloosahatchee July-December I960. Rep. to Off. Nav. River around 6 microgram atoms/ Res., Tech. Rep. 61-3, 109 p. L. . . ." The "method of least squares" is a method usually used for curve A report containing raw hydrographic fitting and can hardly have a bearing data obtained in an area north of Key on calculation of a mean value. Graham West from Cape Sable to Dry Tortugas et al. (1954) obtained average total (July-December, I960). phosphorous values of 12.0 amd 2.6 UNIVERSITY OF MIAMI, MARINE LABORA- ug.at./l. for the Peace and Caloosa- TORY. hatchee Rivers. On page 23 of the Ma- 1962a.. A report of data obtained in Florida rine Laboratory report it is stated that Straits amd off the west coast of Florida out of 22 stations on the Peace River, the (January- June 1961). Rep. to Off. Nav. maxinnum amount of total phosphorus Res., Tech. Rep. 62-4, 200 p. was close to 32 ug.at./l., near phos- phate quarries. It would appear that the A report containing raw hydrographic average derived by "least squares" of data obtained in an area north of Key 31 ug.at./l. is unrealistic. West from Cape Sable to Dry Tortugas About 4 pages of the report list notes (January-June, 1961). on red-tide outbreaks chronologically UNIVERSITY OF MIAMI, MARINE LABORA- from September 1952 through July 1 954. TORY. On September 24, 1953 is the cryptic 1962b. A report of data obtained in Florida note, "Tried CUSO4." Straits and off the west coast of Florida, The report does not give the tech- July-December 1961. Rep. to Off. Nav. nique used for counting G. breve in Res., Tech. Rep. 62-6, 108 p. the samples. However, we believe that their counts are far too high. For in- VEENHUYZEN, J. C.

stance, they stated on p. 73, ". . . With 1879. Communication: Trichodesmium one exception, no discoloration was erythraeum. Natuurk. Tijdschr. Ned. found in which less than 15,000,000 Indie 38:150-151. [Cited from Fish and ." en- Non vidi.] cells/L. occurred. . . This was Cobb. tirely at variance with our experience WALKER, S. T. during the 1957 outbreak, when it was 1884. Fish mortality in the Gulf of Mexico. easily possible to guide spray planes Proc. U.S. Nat. Mus. 6:105-109. onto patches of red tide containing only a few million cells per liter. On November 20, 1880, he sailed

". . . What appeared to be resting south through Boca Ciega Bay and stages of G. brevis were found in salt encountered the first dead fish near water muds. These clearly resembled Bird Key, a little southeast of Pass the forms which appear when G. brevis A'Trilla. On attempting to land on the is partly dessicated. Attempts to con- extreme point of Point Pinellas he was

71 driven back to his boat by the stench dinner." He amd his daughter ate the of thousands of rotting fish upon the oysters and were both sick, but a beach. He mentioned a large variety visitor amd his servant who did not of dead fish and many he could not eat oysters were not affected. identify. He found the same species Two men who owned a fishing smack at Gadsden Point. He saw only a few with a live well stated, "The poisoned dead fish at Tannpa, principally gars water runs in streaks, for often when and catfish. His summary is: three or four smacks are in company "1. The dead fish were most nu- one or two will lose all their fish in a merous on the outside beaches and on few minutes, while others, a short dis- the inside beaches of the outer line of tance off, lose none." keys. The keeper of Egmont Light said first "2. That dead fish were least nu- dead fish appeared on October 17, later nnerous about the mouths of creeks and ducks, and other sea birds. rivers, decreasing gradually as one approached such places. WEBB, JOHN G. "3. That the poisoned water was not 1887. 5. --The mortality of fish in the Gulf diffused generally, but ran in streams of Mexico. U.S. Fish Connm., Bull. of various sizes, as proven by fish 6:11-13. dying in vast numbers instamtly upon reaching such localities. This reference is of no significance. "4. That the fish were killed by a specific poison, as proven by the sick- WENNEKENS, M. P. ness and death of birds which ate of 1959. Water mass properties of the Straits the dead fish. of Florida and related waters. Bull. "5. The fish began dying on the out- Mar. Sci. Gulf Carib. 9(l):l-52. side beaches first, as Mr. Stramd, assistant light-keeper at Egmont, re- Gives a diagram of the general cur- ports them coming up first on the 17th rent system of the eastern Gulf. of October, while Mrs. Hoy observed them first on the 1st or 2d of November, WHITELEGGE, THOMAS. at Little Manatee River. 1891. On the recent discolouration of the "6. The examination of m amy hundred waters of Port Jackson. Rec. Aust. Mus., recently-dead fish revealed no signs Sydney, 1(9): 179-192. of disease. The colors were bright, the flesh firm, and the gills rosy. The A mass mortality caused by Gleno- stomach and intestines appeared dinium rub rum occurred in March 1890, healthy, [p. 106.] at Port Jackson, Australia, during hot "In my haste I have neglected to state weather following heavy rainfall. The that I saw a good many dead birds orgamism occurred in streaks or during the trip. At Tampa, ducks were patches. It caused great devastation dying. I saw dead vultures at Anna Maria annong oysters. Key, and at Passage Key, large flocks of cormorants were sick amd dying. I WILSON, WILLIAM B. also saw the carcasses of terns, gulls, 1955. Laboratory studies of Gymnodinium and frigate birds. The cormorants sat brevis . Address to American Asso- on the beach with their heads under ciation for the Advancement of Science, their wings, and could be approached 14 p. [Mimeographed.] and handled." [p. 106-107.] He appended statements by several In first attempts to culture G. breve , witnesses: orgamisms failed to grow (reproduce ?) "The fish began dying about the last amd reacted as though most media were of October here [Point Pinellas] . . . toxic. Addition of a metal chelator When a school of mullet get into water (EDTA-Na) helped but still only 40 covered by this black scum, they die percent success was attained. The dif- at once. Oysters are affected by this ficulty was chiefly with proper washing also, amd those who eat of them are amd sterilization of the test tubes. made very sick. Good growth resulted from 175 ft.-c. "The oyster saloons here were obliged (foot-candles) for 15 hours per day, but to close, as the oysters came near not from 150 ft.-c. nnaintained continu- killing several people. ously. Eight hours at 175 ft.-c. were "The fish began dying here [Little insufficient. Cloud cover could be a Manatee] about the first of Novem- limiting factor. ber ... A few days after the fish begam G. breve survived 1 hour at 34° C. dying I had a quart of fine oysters for but cytolized at 35° C. When temperature

72 is lowered at 2° per hour, they be- toxic in concentrations of 100 parts per come immobile and cytolize between million. These results indicate that 12° and 14° C, but when lowered at fluorescine dye can be used as a marker 1° per hour they are active until they in field experiments without altering reach 8° to 10° C. the results." [p. 67.] G. breve survive salinities from 24to 40 p.p.t., but become inactive at 20 WILSON, WILLIAM B. p.p.t. or 42 p.p.t. (90 to 100 percent 195 9a. Evaluating toxicity of dissolved sub- of culture). Also see Aldrich and Wilson stainces to microorganisms using di- (1960), alysis membrames. In Galveston Bio- G. breve survived pH from 7.0 to 8.6; logical Laboratory fishery research for growth was best from 7.3 to 8.1. A the year ending June 30, 1959, p. 100- mature culture declining from its peak 102. U.S. Fish Wildl. Serv., Circ. 62. growth frequently reverts to the growth phase if one of several nutrients is To test the effects of copper in the added, namely: B 12. thiamine, biotin, field, known concentrations of Proro- phosphate, ammonia, a group of trace centrum sp. were placed in bags made metals, or soil extract. of dialysis membrane. The bags were If fish autolysate is added to the suspended inside perforated polyethyl- original culture medium, it frequently ene bottles for protection from macro- stinnulates excessive bacterial growth organisms. and the culture dies. In mass cultures "... Between two aind five hours which are cropped and new media added were required for the inside concentra-

the highest concentration of G. breve is tion to approach 0.8 (i. g. at. Cu/l with an normally 4 million per liter, and never outside concentration of 1.6 jj,g.at. Cu/l.

as high as 5 million. Fish autolysate An outside concentration of 0.8 |i g.at. added to a 3.5-million culture raised Cfi/l did not raise the inside concentra- it to 9 1/2 million in 4 days but it fell tion to a comparable amount within 24 to zero in 1 week. hours." [p. 101-102.] The author believed there is a mu- tually beneficial relationship between WILSON, WILLIAM B. G. breve cind "the dominant bacteriad 1959b. Nutritional studies on red tide. In contaminamt of mass cultures." He says Galveston Biological Laboratory fishery G. breve produce carbohydrates which research for the year ending June 30, could account for the increased bac- 1959, p. 72-74, U.S. Fish Wildl, Serv., terial growth, while bacteria produce Circ. 62.

vitamnin Bj2 needed by G. breve . In confirmation of this mutuality of "The minimum calcium content of interests the author had sealed cultures media in which G. breve grew was ap- of G. breve cind G. splendens that were proximately one -sixth the amount of maintaining themselves after 2 amd 2 1/2 standard sea water (salinity - 35. 5%o), years, respectively. or approximately 70 mg/l(l,7 mgat/1). The author could not subculture An increase in calcium to twice the G. breve without adding soil extract amount of open oceain water was not to the medium. Ashed soil extract detrimental as long as the phosphorus would not substitute, amd neither would content was less than 5 |j,g,at/l. If the known chemicals. Soil extract dialyzate calcium content is between 140 and is not beneficial but the dialyzed por- 400 mg/1, cultures have grown very tion remaining is beneficial, indicating well in most concentrations of phos- orgauiic compovmds of high molecular phorus. There was some linnitation of weight. growth if the phosphorus was less tham Of several metals tested, copper was 0.04 p,g.at/l or more thain 100|j.g.at/l. most toxic. In the latter case, the high phosphorus content caused precipitation, as did WILSON, WILLIAM B. excessive calcium. When precipitation 1958. Compounds toxic to red tide orga- occurred, G. breve did not grow. nisms. In Annual report of the Gulf "The results of the calcium -phos- Fishery Investigations for the year phorus experiments indicate that if the ending June 30, 1958. U.S. Fish Wildl, calcium concentration was the same as Serv., p. 66-67. that of normal open ocean water (ap- proximately 400 mg/1), inorgamic phos- Discusses progress on factors af- phorus content should be 0.4(i,g.at. per the to fecting the toxicity of copper to G. breve . liter or greater for medixun

"Tests on the toxicity of fluorescine support good growth of G. breve . Opti- dye to G. breve indicated that it is not mum growth occurred within a rainge of

73 0.4 to 40 [xg.at/l of phosphorus if the Effect of some vitajnins and a group of trace calcium content was between 140 and elements on the growth of G, breve in artificial 400 mg/l. Media with phosphorus con- sea water medium (Numbers represent percent of cultures with each addition in each growth centrations as high as 400 |ig. at/1 sup- category) ported good growth of G^. breve if the calcium concentration was between 140 amd 250 per liter. On the other hand, Additives^ media with calcium concentration as high as 1,000 mg per liter supported good growth if the phosphorus content was between 0.04 and 4.0 |j. g. at/1. The above values for calcium and phosphorus should be revised slightly upward (less than 0.01 percent) because of the occur- rence of these elements as contaminants in the other components of the medium. [p. 72.] " Vitamins . Three water soluble vita- mins, thiamine (Bj), biotin (B3) and cobalamin (B12 )> have been used con- sistently in artificial sea water media for bacteria-free G. breve . Results of earlier experiments indicated that G. breve would not grow in these media unless these vitamins were added. In addition, thiamine was the most effec- tive of the three vitannins, but a com- bination of thiamine and biotin supported better growth than thiamine alone. Dur- ing the past year, a series of nine experiments was conducted to define more clearly the role of these vitamins in the growth of G. breve, [p. 72-73.] "Results of these experiments are sinnilar in many ways to the results of previous work in that thiamine was the most active of the three vitamins and a connbination of thiamine and biotin was better thain thiamine alone. How- ever, these experiments indicate that the inclusion of cobalamin along with the thiamine and biotin further im- proved growth. In the earlier experi- ments, we had omitted cobalamin in ten serial subcultures without diminution of growth. In recent experiments, omis- sion of cobalamin has resulted in lesser growth. To ascertain the source of the apparent discrepancy in these results we will initiate growth experiments using cobalamin Eind the other vitamins from more tham one supply house, [p. 73.] "The need for thiamine is pronounced in the experiments conducted to date. There is a possibility that similar or- ganic compounds will substitute for thiannine and we plan to conduct experi- ments to determine if such is the case. If not, assays of this vitamin in the field may be valuable for forecasting red tides, [p. 73-74,] " Trace elennents . In earlier experi- ments we were unable to grow G. breve in artificial sea water nnedia unless a complexing substances to prepare a C. E. Dawson and Henry Hildebrand air south of the Rio Grande suitable sea water medium for G. breve , checked by in most instances. Therefore, the oc- and observed dead fish at several points. currence of natural metal chelators or One water sample taken 36 miles south metal complexing substances may be of the Rio Grande contained over 22,000 necessary for red tides to develop." per ml. At least 120 miles of the fish [p. 74.] Tamaulipas coast was affected; were still dying on September 27, 1955. WILSON, WILLIAM B. I960. Red tide investigation. In Galveston WILSON, W. B., and SAMMY RAY. Biological Laboratory fishery research 1958. Nutrition of red tide organisms. In for the year ending June 30, I960, p. 39. Annual report of the Gulf Fishery In- U.S. Fish Wildl. Serv., Circ. 92. vestigations for the year ending June 30. 1958, U.S. Fish Wildl. Serv., p. 62- Summary of fiscal year progress 65. given in detail in other reports. Discusses the development and gives the formula for an artificial sea water WILLIAM B., and WILSON, ALBERT COLLIER. medium for growing bacteria-free cul- 1955. Prelinninary notes on the culturing of tures of G. brevis . Gymnodiniunn brevis Davis. Science "G. breve requires phosphorus for 121(3142):394-395. . growth in this artificial medium, but the absolute amount needed for growth ". . Titanium and zirconium were . has not been determined because the added to the mixture, although they are major salts contain phosphorus as an present in not normally measurable impurity. Cultures grew well without in water, because these amounts sea adding phosphorus until we recrystal- metals were found abundantly in water lized the sodium chloride to increase of G. brevis blooms, [p. 394.] its purity. Increasing the phosphorus additions above the minimum amoiant necessary for growth does not increase the number of organisms. The addition "Laboratory cultures of G. brevis of 0.1 microgram atom of phosphate have attained homogeneous concentra- phosphorus per liter (0.1 l^g.at. P/L) tions exceeding 2 million cells per is sufficient to support good growth. concentration is far below liter. This Chemical analyses of the medium for the highest report; however, the values total phosphorus with this amount added cannot be compared. Among other rea- indicate that it contains between 0.1 sons, G. brevis in cultures concentrate and 1.0 at. P/L. Therefore, the disperse by fig. to form masses that we amount of phosphorus required for making counts. similar shaking before A maximum growth in this medium is concentrate, but on a larger tendency to apparently about 1.0 (ig.at. P/L or scale, may be expected in nature. Counts ." less. . . [p. 64.] of a sample from such a concentration

would be high. . . ." [p. 395.] WOOD, E. J. F. 1954. Dinoflagellates in the Australian Re- WILSON, W. B., and S. M. RAY. gion. Aust. J. Mar. Freshwater Res. 1956. The occurrence of Gymnodinium 5{2):171-351. brevis in the western Gulf of Mexico, Ecology 37(2):388. Lists and describes each species. Diagrams of species are outline draw- From 1953 to 1955 sea-water samples ings. No Gymnodinium listed. He specu- were collected at Galveston and along lated on correlation of Gonyaulax digi- the Gulf coast between Galveston and tale with the new moon, but the evidence Florida, but no G. brevis were ob- presented is not conclusive. served. In September 1955, 12 samples in WOOD, E. J. F. about 20 square miles from the mouth 1963. Dinoflagellates in the Australian Re- of the Rio Grande to 10 miles N. of gion. II. Recent collections. Common- Port Isabel all contained from 50 to 500 wealth Sci. Ind. Res. Org., Div. Fish. G. brevis per ml. One sample near Oceanogr., Tech. Pap. 14, 55 p. Port Isabel in the Laguna Madre con- lists tained no G. breve . Dead fish were on Describes 16 new species and Gulf beaches from Rio Grande to 17 over 190 species found since 1954.

miles N. of Port Isabel. Lists 30 species of Gymnodinium .

75 WOODCOCK, ALFRED H. SUBJECT INDEX TO SELECTED TOPICS 1948. Note concerning human respiratory irritation associated with high concen- Acartia trations of plankton and mass mortality abundant in G. breve blooms of nnarine organisnns. J. Mar. Res. Univ. Miami, Mar. Lab., 1954 7(l):56-62. Acartia tonsa water discolored by red -tide water, when Shows that the Hutton, 1960 sprayed into the nostrils or when breathed as an aerosol, caused coughing Airplame and a burning sensation in the nose aind use in red tide research throat. Such aerosols are formed when Ingle et al., 1959 waves break on the beach. Breathing through a 2-cm. pad of Amphidinium fusiforme absorbent cotton prevented irritation. Martin and Nelson, 1929 Unconcentrated "red water" contain- Pomeroy et al., 1956 ing 56 million G. breve per liter re- tained its irritating qualities unim- Annphidinium klebsii (?) sifter for several weeks. paired storage McLaughlin and Provasoli, 1957 The irritant passed through a fine bacteria filter (openings, 1 to 1.5 Amphidinium rhyncocephalus microns). McLaughlin and Provasoli, 1957

WOODCOCK, A. H. Aniline dye in weed control 1955. Bursting bubbles and air pollution. Eicher, 1947

Sewage Ind. Wastes 27(10): 1 189- 1 1 92. Aulacodiscus kittoni Becking et al., 1927 "At relative humidities commonly found over the sea, droplets of sea Bacteria of this size [2-200 fi.] will evap- water chromogenic, toxicity of orate quickly to form nuclei of about Bein, 1954, 1955 1 to 45 in diameter. Such nuclei fall fx Buck et al., 1963 slowly (0.01 to 13 cm. per sec.), aind Ray and Wilson, 1957 sind winds cause atmospheric turbulence as detoxicating agents carried mciny miles and them to be Ray and Wilson, 1957 hundreds of feet into the air .... Shilo cind Aschner, 1953 [p. 1189.] as vitamin Bi2 producers Wilson, 1955

Bibliography Brunei et al., 1950 "Microscopic examination of the aero- Fish and Cobb, 1954 revealed that sols from 'red water' Fritsch, 1935 their surface tension was much re- Hayes cind Austin, 1951 the surface tension duced, compared to Hutton, 1956 of droplets of relatively clear sea Provasoli, 1958 water, and that some of the droplets contained parts of plankton orga- Calcium ." nisms. . . [p. 1191.] effect on growth of G. breve Wilson, 1959b phosphorus-calcium relation effect on ACKNOWLEDGMENTS growth of G. breve Wilson, 1959b Mciny authors and publishers have permitted us to quote from their articles. Annong the Calcium hypochlorite periodicals are Advances in Food Research, use in red-tide control Jour- Americcui Geophysical Union, American Galtsoff, 1948 nal of Science, Botainiceil Gazette, Economic Geology, Limnology and Oceanography, Carbohydrates Science, Scientific American, The Americam methodology of determining Naturalist, The Geological Society of America, Eldred et al., 1964 Inc., and The Journal of Protozoology. Lewis and Rcikestraw, 1955

76 Ceratia Dialysis membranes Nordli, 1953 bioassays using Wilson, 1959a Ceratium furca Hutton, 1956 Discolored water Ryther, 1955 causes of Becking et al., 1927 Ceratium fusus Hirasaka, 1922 Ryther, 1955 Hutton, 1960

Ceratium hirundinella Evadne effect of copper sulphate on ingestion of G. breve by Galtsoff, 1948 Galtsoff, 1948 Cochlodinium as chain -forming Exuviella baltica Lackey aind Hynes, 1955 Numann, 1957 Collecting gear Anderson, 1951 Flavobacterium piscicida Arnold, 1958 Bein, 1954, 1955 King, Joseph, 1950 Buck et al., 1963 Lackey cind Hynes, 1955 Ray and Wilson, 1957 description of Convergence mechanism Bein, 1954 Chew, 1955a Buck et al., 1963 (renamed Pseudomonas ) Ewing, 1950 Ingle et al., 1959 Fluorescine dye Langmuir, 1938 non -toxic to G. breve Pomeroy et al., 1956 Anon., 1956 Ryther, 1955 Wilson, 1955 Stommel, 1949 Copper Gonyaulax sensitivity to estuarine fauna fish kills by Aldrich, 1958a Bonnot and Phillips, 1938 Copper ore Connell and Cross, 1950 control for red tide Gates, 1958 Marvin, 1958, 1959 Gates and Wilson, I960 Marvin et al., 1961 Howell, 1953 Wilson, 1959a Kofoid, 1911 Mitsukuri, 1905 Copper sulphate Nishikawa, 1901 control for red tide Torrey, 1902 Cahn, 1950 Collier, 1955 Gonyaulax Crance, 1963 occurrence Fisher, 1956 Davis, 1955 Galtsoff, 1948 Hale, 1950 Gonyaulax monilata Ingle et al., 1959 description of Rounsefell, 1958 Howell, 1953 Rounsefell and Evans, 1958 Sater, 1954 Shila et al., 1954 Gonyaulax polyedra Haxo cind Sweeney, 1955 Coscinodiscus Kofoid, 1911 Hutton, 1956 Sweeney cind Hastings, 1958 Culture - see Physiology Gymnodinium Culture techniques gelatinous envelope of isolation for pure cultures Martin and Nelson, 1929 Brunei et al., 1950 Droop, 1954 McLaughlin, 1958 Gynnnodinium adriaticum of Soli, 1964 development from zooxanthellae Spencer, 1952 Hutner and McLaughlin, 1958

77 Gymnodinium breve Gyrodinium californicum bioassays of Ryther, 1955 Ray and Wilson, 1957 Starr, 1958 Heterotrophy density required to detect discoloration in photosynthetic dinoflagellates Lackey and Hynes, 1955 Provasoli and McLaughlin, 1963 description of Davis, 1948 Histological techniques amd Hynes, 1955 Lackey G. breve dominance in samples of Aldrich, 1958a Davis, 1955 Finucane, 1959b cleavage of Arbacia egg of effect on Lackey and Hynes, 1955 Cornman, 1947 Martin and Nelson, 1929 enumeration techniques for Lackey and Hynes, 1955 Hurriccines Mar. Lab., 1954 Univ. Miami, Dragovich and May, 1961 life stages of Feinstein, 1956 Lackey and Hynes, 1955 Gunter et al., 1948 Lab., 1954 Univ. Miami, Mar. Ingersoll, 1882 occurrence at Trinidad of Lackey, 1956 Hydrography occurrence in the Western Gulf of Florida, west coast of Burr, 1945 Chew, 1955a, 1955b Gunter, 1952 Eldred et al., 1964 Lund, 1935 Hela et al., 1955 Wilson and Ray, 1956 Koczy et al., I960 offshore survival of Marvin, 1955 Dragovich, I960 Tabb et al., 1959, 1962 Dragovich and May, 1961 Gulf of Mexico Finucane, 1960 Collier et al., 1958a rate of division of Graham et al., 1954 Collier, 1955 Ichiye, 1962 Lackey and Hynes, 1955 Salsman and Tolbert, 1963 Wilson, 1955 Univ. Miami, Mar. Lab. 196la, 196lb, sampling of, extreme patchiness of distribu- 1962a, 1962b tion in Wennekens, 1959 Finucane, 1958 Naples area seasonal occurrence of Dragovich, 1963 Finucane, 1964 vertical diurnal movement of Hydrology Finucane, 1958 Tampa Bay viscosity of water caused by Dragovich and Kelly, 1964 Galtsoff, 1948 Dragovich and May, 196l Gunter at al., 1947 May, 1964 Gymnodinium flavum Olson, 1953 occurrence in California of Kofoid, 1911 Isochrysis galbana Lackey and Clendenning, 1963 McLaughlin, 1958 mikinnoti Gymnodinium Labidocera aestiva Matoda, 1944 water discolored by Gymnodinium sanguineum Hutton, 1960 Hirasaka, 1922 Kofoid and Swezy, 1921 Larval forms killed by red tide ymnodinium simplex G Matoda, 1944 bacteria-free cultures of King, Gladys, 1950 Mesodinium G ymnodinium splendens Clemens, 1935 Pomeroy et al., 1956 Hart, 1934 bioassay of Slobodkin, 1953 Collier, 1954 use in experiments of Metabolites Collier, 1954 ecological effect of Sweeney, 1951, 1954 Lucas, 1947, 1949 78 Metals PH role in G. breve metabolism of changes of as control measure Hutner et al 950 Galtsoff, 1948 Starr, 1958 Monochrysis luthe ri McLaughlin, 1958 Phosphorus calcium-phosphorus relation, effect on Mycrocystis growth of G. breve of Crance, 1963 Wilson, 1959b concentration by Skujaella Nitrogen Graham et al., 1954 effect on red tide outbreaks of concentration in river discharges of Dragovich, 1960 Dragovich, 1959, 1960b Gilson, 1937 Graham et al., 1954 Lackey and Hynes, 1955 Odum, 1953 Specht, 1950 Noctiluca concentration in West Florida rivers of mortality caused by Alafia River Brongersma-Sanders, 1948 Dragovich, 1959, 1960b Dragovich and May, 1962 Caloosahatchee River Noctiluca miliaris Dragovich and May, 1962 Bhimachar and George, 1950 Univ. Miami, Mar. Lab., 1954 Pratje, 1921 Hillsborough, Little Mematee, and Manatee Rivers Nutrition - see Physiology Dragovich, 1959 Dragovich and May, 1962 Peace River Offatts Bayou Dragovich and May, 1962 mortality of fish in Graham et al., 1954 Connell amd Cross, 1950 Specht, 1950 Gates and Wilson, 1960 Univ. Miami, Mar. Lab., 1954 effect on red tide outbreaks of Orinoco River Bein, 1957 relation to red tide outbreak of Chew, 1953 Lackey, 1956 Graham et al., 1954 Lackey and Hynes, 1955 Oysters methodology of determining killed by red tide Robinson and Thompson, 1948 Anon., 1934 (in Japan) minimum levels for blooms Gaits off, 1948 Lackey and Sawyer, 1945 Jefferson, 1879 occurrence in particulate form Mitsukuri, 1905 Graham et al., 1954 Miyajima, 1934 origin of Rathbun, 1895 Chew, 1953, 1955b U.S. Fish Wildl. Serv., 1961 Dragovich, 1959 Whitelegge, 1891 Graham et al., 1954 red colored fronn red tide Lasker and Smith, 1954 Clemens, 1935 Phelps and Barry, 1950 Nelson, 1948 Smith, 1949 Pomeroy et al., 1956 Specht, 1950

Peridiniales Physiology of G. breve Graham, 1942 carbohydrates, effect on growth of Aldrich, I960 Peridiniunn Collier, 1958b Mead, 1898 concentrations in laboratory cultures, com- parison with natural Peridinium sainguineum Wilson and Collier, 1955 Carter, 1858 copper, toxicity of Aldrich, 1959 Peridinium triquetum fish autolysate, effect of Braarud and Pappas, 1951 Collier, 1955 Ryther, 1955 Wilson, 1955

79 Physiology of G. breve --Continued Lucas, 1947, 1949 light, Margalef, 1963 intensity of Menon, 1945 Aldrich, 1960 Rice, 1954 Brongersma-Sanders, 1957 Ryther, 1955 Lackey and Hynes, 1955 volumes of Wilson, 1955 Arnold, 1958 polarized Polychaetes Aldrich, 1960 immunity to G. veneficum toxin of wave length Ballantine and Abbott, 1957 Aldrich, 1960 nutrition, autotrophic Polykrikos Aldrich, I960, 1962 harmless to pearl oysters pH, effect of Galtsoff, 1948 Aldrich, 1959 McLaughlin, 1956 Predators of dinoflagellates Wilson, 1955 Galtsoff, 1948 river water from various sources, effect Hutner and McLaughlin, 1958 on growth of Torrey, 1902 Aldrich, 1961 sailinity, effect of Productivity Aldrich, i960 comparison of Gulf to other areas of Aldrich and Wilson, I960 Anderson, 1951 Dragovich, 1963 Wilson, 1955 P rorocentrum nniceins synthetic media Allen, 1933 King, Gladys, 1950 Braarud and Rossavik, 1951 Ray and Wilson, 1957 Numann, 1957 Wilson, 1955 Sweeney, 1951 Wilson and Collier, 1955

Wilson and Ray, 1958 P rorocentrum sp . temperature, effect of use as test organism for toxicity of copper Aldrich, 1959, 1960 Wilson, 1959a Barker, 1935 Dragovich, 1963 Prymnesiunn parvum Dragovich and May, 1961 control of in ponds Finucane, 1960 Shila et al., 1954 Wilson, 1955 red tide caused by trace elements, effect on growth of OtterstrjZ^m and Nielsen, 1939 Wilson, 1955 thiamine requirements of tyrosine, effect on growth of Rotberg, 1958 Collier, 1958b toxicity of, effect of pH on vitamins, effect on growth of McLaughlin, 1956 Haxo and Sweeney, 1955 Shilo and Aschner, 1953 McLaughlin and Provasoli, 1957 piscicida Provasoli, 1958 Pseudomonas ( Flavobacterium ) Wilson, 1959b revision of Flavobacterium Buck et al., 1963 Plagicampa marina bacteria-free cultures of Rainfall King, Gladys, 1950 effect on outbreaks of Burr, 1945 Plankton Collier, 1954 occurring in Florida Lund, 1935 Davis, 1950 Menon, 1945 Davis and Williams, 1950 Numann, 1957 Dragovich, 1961, 1963 Slobodkin, 1953 King, Joseph, 1950 Whitelegge, 1891 succession of species of Bein, 1957 Red tide Brongersma-Sanders, 1957 aerosols of Finucane, 1959a Gunter et al., 1947 Hart, 1942 Ingle, 1954 Hornell and Nayudu, 1923 Lund, 1935 Hutchinson, 1944 Woodcock, 1948. 1955

80 Red tide --Continued occurrence on Campeche Banks control of Graham, 1954 by chemicals occurrence in Iceland Anon., 1934 (Ferric chloride, chlorine) Paulson, 1934 Cahn, 1950 occurrence in India Collier, 1955 Bhimachar and George, 1950 Galtsoff, 1948 Carter, 1858 Hutner and McLaughlin, 1958 Denison, 1862 Lackey and Hynes, 1955 (Charcoal) Hornell, 1917 Marvin, 1958, 1959, 1960 Hornell and Nayudu, 1923 Marvin and Proctor, 1964 Jouan, 1875 McLaughlin, 1958 (Ammonia) Menon, 1945 Odum et al., 1955 occurrence in Japan Rounsefell, 1958 Mitsukuri, 1905 Rounsefell and Evans, 1958 Nishikawa, 1901 Univ. Miami, Mar. Lab., 1954 occurrence in Lower California by fish culture in bays Stohler, I960 Univ. Miami, Mar. Lab., 1954 Streets, 1878 by high-frequency radio waves occurrence in Narragansett Bay Lackey and Hynes, 1955 Mead, 1898 by seining dead fish Sherwood and Edwards, 1902 Univ. Miami, Mar. Lab., 1954 occurrence in Sicily effect on birds of Forti, 1933 Galtsoff, 1948 occurrence in Southwest Africa Taylor, 1917 a and b Brongersma-Sanders, 1945, 1948, 1957 Walker, 1884 Classen, 1930 effect on edible oysters of Hart, 1934 Cahn, 1949, 1950 Kaiser, 1930 Graham, 1954 occurrence in Tunis Walker, 1884 Cabasso and Roussel, 1942 effect on pearl oysters of occurrence in 1844 Anon., 1934 IngersoU, 1882 Cahn, 1949, 1950 Taylor, 1917 a and b Mitsukuri, 1905 occurrence in 1854 Miyajima, 1934 Ingersoll, 1882 Nishikawa, 1901 Taylor, 1917 a and b Florida data on occurrence in 1865 Dragovich et al., 1963 Glazier, 1882 Dragovich et al., 1961 occurrence in 1878 Finucane and Dragovich, 1959 Glazier, 1882 Goodell and Gorsline, 1961 Ingersoll, 1882 Odum et al., 1955 Jefferson, 1879 numbers of fish killed by Jefferson et al., 1879 Finucane et al., 1964 Moore, 1882 Gunter et al., 1948 Taylor, 1917 a and b Lackey and Hynes, 1955 occurrence in 1879 Smith, 1954 Ingersoll, 1882 occurrence in Aegean Sea occurrence in 1880 Niimann, 1957 Glazier, 1882 occurrence in Algeria Ingersoll, 1882 Hollande and Enjumet, 1958 Moore, 1882 occurrence in Angola Taylor, 1917 a and b Niimann, 1957 Walker, 1884 occurrence in Arabian Sea occurrence in 1882 Lees, 1937 Anon., 1883 occurrence in Australia Taylor, 1917 a and b Whitelegge, 1891 occurrence in 1883 occurrence in California Gunter, 1947 Allen, 1933, 1935, 1943, 1946 Taylor, 1917 a and b Bonnot and Phillips, 1938 occurrence in 1884 Kofoid, 1911 Gunter, 1947 Lackey and Clendenning, 1963 occurrence in 1885 Sommer and Clark, 1946 Canova, 1885 Torrey, 1902 Glennan, 1887

81 Red tide --Continued major river drainages in relation to occurrence in 1908 general Taylor, 1917 a and b Finucane, 1958 occurrence in 1916 Lackey, 1956 Taylor, 1917 a and b Long, 1953 occurrence in 1935 Univ. Miami, Mar. Lab., 1954 Lund, 1935 (in Texas) Apalachicola River occurrence in 1946 Chew, 1955a, 1956 Galtsoff, 1948 Caloosahatchee River Gunter et al., 1947 Collier, 1953a Gunter et al., 1948 Peace River occurrence in 1947 Univ. Miami, Mar. Lab., 1954 Davis, 1948 movement of Galtsoff, 1948 Chew, 1955b Gunter et al., 1947 Hela et al., 1955 Gunter et al., 1948 Taylor, 1917 a and b occurrence in 1948 Univ. Miami, Mar. Lab., 1954 Gunter, 1952 (in Texas) movement of fish killed by occurrence in 1949 Hela et al., 1955 Feinstein, 1956 occurrence in streaks of occurrence in 1952 Anon., 1883, 1956 Chew, 1953 Bonnot and Phillips, 1938 Collier, 1953a Hornell and Nayudu, 1923 Lackey and Hynes, 1955 Niimann, 1957 Smith, 1957 Pomeroy et al., 1956 Univ. Miami, Mar. Lab., 1954 Taylor, 1917 a and b occurrence in 1953 Torrey, 1902 Chew, 1953 Whitelegge, 1891 Collier, 1954 origination in passes of Lackey and Hynes, 1955 Hela, 1955 Univ. Miami, Mar. Lab., 1954 Ingle et al., 1959 occurrence in 1954 Univ. Mianni, Mar. Lab., 1954 Finucane, 1964 phosphorus in relation to Lackey and Hynes, 1955 Bein, 1957 Univ. Miami, Mar. Lab., 1954 Chew, 1953 occurrence in 1955 Graham et al., 1954 Finucane, 1964 Ketchum and Keen, 1948 Hutton, 1956 Lackey and Hynes, 1955 occurrence in 1956 Odum, 195 3 FinucEine, 1964 Univ. Miami, Mar. Lab., 1954 occurrence in 1957 prediction of Finucane, 1964 Chew, 1956 occurrence in 1958 Feinstein, 1956 Finucane, 1958 Smith, 1957 occurrence in 1959 rainfall in relation to Finucane, 1960 Burr, 1945 occurrence in 1960 Collier, 1954 Finucane, 1960, 1961 Dragovich and Kelly, 1964 occurrence in 1963 Feinstein, 1956 Ingle and Sykes, 1964 Hornell and Nayudu, 1923 Finucame et al., 1964 Lund, 1935 outbreaks of Niimann, 1957 decaying fish sustain salinity in relation to Brongersma-Sanders, 1957 Chew, 1955c Collier, 1954, 1955. 1958b Dragovich and Kelly, 1964 far offshore Finucane, 1960 Finucane, 1964 Nordli, 1953 Hutton, 1960 Ryther, 1955 Lackey and Hynes, 1955 storms in relation to historical intensity of Feinstein, 1956 Feinstein, 1956 Ingersoll, 1882 Feinstein et al., 1955 stream discharge in relation to lunar phase in relation to Chew, 1955a, 1956 Feinstein et al., 1955 Feinstein, 1956 Univ. Miami, Mar. Lab., 1954 Wood, 1954 82 Red tide --Continued Seawater submarine springs in relation to biochemistry of Graham et al., 1954 Armstrong and Boalch, 1961 Univ. Miami, Mar. Lab., 1954 Collier, 1958b sulfides in relation to Pomeroy et al., 1956 Lackey and Hynes, 1955 McKee and Wolf, 1963 Sewage sunshine in relation to algal bloonns caused by Brongersma-Sanders, 1957 Braarud, 1945 Cahn, 1950 Brongersma-Sanders, 1957 Clemens, 1935 Connell and Cross, 1950 Conover, 1954 Lackey, 1958 Hornell, 1917 Odum, 1953 tennperature in relation to Phelps and Barry, 1950 Aldrich, 1959 Barker, 1935 Shellfish toxicity Brongersma-Sanders, 1957 relation of red tide to Gunter et al., 1948 Abbott and Ballantine, 1957 Nordli, 1953 Eldred et al., 1964 Ryther, 1955 Gulf Coast ShellfishSanit. Res. Cent., 1964 tidal phases in relation to La Cossitt, 1954 Chew, 1955c Lackey and Hynes, 1955 upwelling in relation to McFarren, 1963 Brongersma-Sanders, 1945, 1948, 1957 McFarren et al., 1957, I960 Graham et al., 1954 Sommer et al., 1937 Gunther, 1936 Walker, 1884 water mass theory in relation to Anderson, 1952 Skujaella erythraeum Chew, 1953, 1956 donninant in Tampa Bay Conover, 1954 Finucane, 1960 Hela, 1955

Kierstead and Slobodkin, 1953 Skujaella ( Trichodesmium ) erythraeum Slobodkin, 1953 Veenhuyzen, 1879 Univ. Mianni, Mar. Lab., 1954 occurrence and effects in India worldwide occurrence of Chidambaram and Anny, 1944 Ballantine and Abbott, 1957 phosphorus concentration by CoUingswood, 1868 Graham et al., 1954 Dareste, 1855 Lackey and Hynes, 1955 Hutner and McLaughlin, 1958

Nightingale, 1936 Skujaella ( Trichodesmium ) thiebauti reported from Campeche Banks mistaken for red tide Graham, 1954 Hutton, 1960 reported from Dry Tortugas water discolored by Chew, 1953 Hutton, 1960 Gunter et al., 1948 Jefferson et al., 1879 Soil extract species killed by use in dinoflagellate culture Finucane et al., 1964 Sweeney, 1951 Galtsoff, 1948 Wilson, 1955 Gunter et al., 1948 Lackey and Hynes, 1955 Tampa Bay Moe, 1964 phosphorus content of Sonnmer and Clark, 1946 Dragovich, 1959 Taylor, 1917 a and b relation of volume of Bay to river discharge research program Odum et al., 1955 Sater, 1954 Olson, 1953 Smith, 1954 Taxonomy, general "Red Tide Index Number" Bursa, 1963 (Arctic Alaska) Chew, 1956 Graham and Bronikovsky, 1944 Graham, 1954 (Peridiniales) Rhythmic cell division Schiller, 1933, 1937 Sweeney and Hastings, 1958 Wood, 1954, 1963 (Australia)

83 Tides Ultra-violet absorption of sea water effect on outbreaks Armstrong and Boalch, 1961 Chew, 1955c May, 1964

Vertical migration Titanium Hasle, 1950, 1954 in red tide bloonn Peters, 1929 Collier, 1953b Pomeroy et al., 1956 Ryther, 1955 Toxicity of red-tide killed fish Vitamin B 12 Hornell, 1917 distribution in marine waters of Lackey and Hynes, 1955 Burkholder and Burkholder, 1956 Taylor, 1917 a and b Cowey, 1956 Walker, 1884 Eldred et al., 1964 bioassays of, for G. breve Starr, 1956 Collier, 1954 indicator orgainisms of Ray and Wilson, 1957 Eldred et al., 1964 Hutner and McLaughlin, 1958 Toxin Ryther and Guillard, 1962 action of dinoflagellate Covell and Whedon, 1937 Gymnodinium splendens requirement of Hutner and McLaughlin, 1958 Sweeney, 1954 bioassays of Wind

( Flavobacte rium piscicida Pseudomonas ) effect on outbreaks of Bein, 1954 Becking et al., 1927 Buck et al., 1963 Bhimachar and George, 1950 Ray and Wilson, 1957 Clemens, 1935 Gymnodinium veneficum Hornell, 1917 Abbott and Ballantine, 1957 Hornell and Nayudu, 1923 Ballantine and Abbott, 1957 Lackey and Hynes, 1955 attributes of Gymnodinium breve , Pomeroy et al., 1956 Starr, 1958 action of G. veneficum Zirconium Abbott and Ballantine, 1957 in red tide bloom Ballantine and Abbott, 1957 Collier, 1953b

84 Appendix table 1. --Monthly rainfall (inches) for south Florida, 1910-60

Month

01726 5 WHSE Created in 1849, the Department of the Interior— a depart- ment of conservation—is concerned with the management, conservation, and development of the Nation's water, fish, wildlife, mineral, forest, and park and recreational re-

sources. It also has major responsibilities for Indian and Territorial affairs. As the Nation's principal conservation agency, the De- partment works to assure that nonrenewable resources are developed and used wisely, that park and recreational re- sources are conserved for the future, and that renewable resources make their full contribution to the progress, pros- perity, and security of the United States—now and in the future.

UNITED STATES POSTAGE AND FEES PAID DEPARTMENT OF THE INTERIOR U.S. DEPARTMENT OF THE INTERIOR FISH AND WILDLIFE SERVICE BUREAU OF COMMERCUL FISHERIES WASHINGTOX, D.C. 20240

OFFICIAI. BUSINESS

Return this sheet to above address, if you do NOT wish to receive this material Q^, or if change of address is needed [31 (indicate change including ZIP Code).

^ Librarian SSR 7 Marine Biological Lab., Wools Hole, Mass. 02543