LEVEhCIDPatterns of Bilwnuineseence In the Ocens

R. V. Lyw04 III

doomom SckuMp*im~

Apri 23, 19"1

MMECTE

S815l' 1 198

_ _ _ _ _ Q 112

REPOT DCUMNTATON AGEREAD INSTRUCTIONS

REPOT DOUMETATIN PAE 1EFORE COMPLETING FOFkM

(/9, 8E~mZ~hWaGOVT ACCESSION NO0 3. tCI01ENY'S CATAL.0G NUMBER

4 6 I YEOFRPRT & PECRIOO C0O/ERID

PATTERNS OF BIOIN T I4E SU N Final reltnnNRprbe

P, 4. I ERVORMING ORO, REPORT NUMBER

7. AUTHOR(s) 0. CONTRACT ONcRAHT NUMBER(*)

R.V. Lych, Navy"Sikqsce AssistanceLyncd~flPrograp",6EV-7-80

9- PERFORMING OGNZTNNAHEAN ADDRESS 10. PROORAM ELEMENT, PROJECT, TASK

ORANZA~O NMEAREA & WORK UN IT NUMBERS

Naval Research LaboratoryWashington, DC 20375 (~j~~~L // 43-1 121A-1

11 CONTROLLING OFFICE NAME AND ADD.RESS 12. REPORT OA;T

E

Naval Research Laboratory Apri 23MBE 1981GEWashington, DC 20375 4 SNMERO AE

-V1.j~TbiINZAG~EN-CY 'NAME &APORESS(l( different froal controllin'g Office) IS. SECURITY t:L ASS. (of It.reot

jJNCLASSUPIEDIS&.ODCLASSIFICATIONI DOWNGRADING

SCI4EOULE

16. DISTRIBUTION STATEMENT (of thA. Report)

Approvred for public release; distribution unlimited.

97. DISTRIBUTION STATEMENT (of the abstrae entereod In Block 20, It dlifferent from Popoft)

IS. SUPPLEMENTARY NOTES

It. KEY WOROS (Conlinuu an evotOOoIo sieIt noroO*O*T 4nd Idenftl4 by hlock ntumbe:)

Bioluminescence distributionArabian SeaPhosphorescent displays

to. AT (Cntine onreverse side it noeoosery and Identify by, block nustbot)

Bioluminescence o,~curs throughout the world's oceans at all times of year. However, the 'possibilities of encountering it vary according to location and season. Maps indicating thesepossibilities are presented in this report, along with a discussion of tP~efr reliability and thestatistical problems involved In their construction and interpretation.

DO ~~S1473 EDITION OF I NOV 61 IS OBSOLETESN0 10 2-014 6601

SECURITY CLASSIFICATION Of THIS PAGE (fte"f Date BItrdoO)

CONTENTS

INTRODUCTION .............................................................................. 1

METHODS ....................................................................................... 2

RESULTS ............................................................................ ............. 2

DISCUSSION AND CONCLUSIONS........................................................... 3

Atlantic Ocean.................................................................................. 3Gulf of Mexico and Caribbean Sea ........................................................... 3Mediterranean and Black Seas................................................................. 3Pacific Ocean.................................................................................... 13Indian Ocean ................................................................................... 13-

STATISTICAL CONSIDERATIONS ........................................................... 29

CORRELATIONS................................................................................ 30 .

ACKNOWLEDGMENTS........................................................................ 30

REFERENCES .......... ........................................................................ 31

AliiACC85SOfli nT

III

PATTERNS OF BIOLUMINESCENCE IN THE OCEANS! • ,iINTRODUCTION

Bioluminescence, or the emission of light by living organisms, is a world-wide phenomenon. ? iscaused by many creatures, the best-known of which is the firefly. However, there are a greater numberand variety of luminous marine organisms than of luminous terrestrial or fresh-water organisms. Con-sequently, light in the sea is often seen.

Luminous marine species occur throughout the animal kingdom from the one-celled iadiolariansto fish and squid. In the plant kingdom, only certain dinoflagellates are luminous. Some marine bac-teria also luminesce. In some areas of the oceans, luminous species are the dominant forms of life.For example, below the photic zone of the ocean it has been estimated that 70% of the species and 90%of the individual organisms are luminous [1). Even near the surface luminous species are frequentamong the plankton. Furthermore many luminous midwater fish, most notably myctophids, and otherorganisms such as euphausiid crustaceans and squid migrate mertically to the surface at night to feed.

With such a diversity of luminous organisms, there are great variations in luminescence and atleast five chemical mechanisms for producing light have been identified. Means of physiological controlare as varied as the organismsw themselves. The biological roles of luminescence are largely unknown,but several have been determined for a limited number of orgrnisms. The intensity, kinetics and dura-tion of the flash, patterns of flashing, and peak emission wavelength differ for each species. In fact,these characteristics are so species-dependent that it might be possible to identify the emitting organ-isms by measuring these five parameters.

F-w luminous marine organisms have been studied in detail; consequently, little is known aboutthe bioluminescent behavior and characteristics of the vast majority. The best source of information isthe book Bioluminescence by E.N. Harvey [2]. This work has been updated for marine species in gen-eral by Tett and Kelly [3), Herring [41, and Kelly and Tett [5). In addition, a bibliography withabstracts of individual papers on aspects of bioluminescence of naval interest, concentrating on work N

done since 1952, has been compiled [61 and a technical assessment of the bibliography prepared 17). -

Most observations of bioluminescence at sea have simply been recorded as the presence of lightand have not been concerned with the identification *of the causative organisms. Displays haveappeared to be both spontaneous, with no identifiable source of stimulation, and as the result of stimu-lation by mechanical means. Most luminous species can be expected to have only a limited geographi-cal range but some higher groups (notahly euphausiids, copepods, and dinoflagellates) appear to beuniversally distributed in the oceans [11.

Several previous studies based primarily on visual sightings have been made on the distributionand occurrence of marine bioluminescence. The earliest general studies were those of Smith [8,91, whoexamined the Meteorological Logs and Records of the Voluntary Observing Fleet from 1920-1930.Regional studies were made of the Okhotsk Sea [101, the Arabian Sea [1I,12J, the Atlantic Ocean 1131,and the North Sea [141. More recent general studies have beer those of Turner [15,161, and Staples[171. Turner [151 made the first attempt at statistically interpreting the sighting reports to identify areasof high bioh, minescence by considering the shipping density as well as the number of sightings. LynchW1] included in his report not only sightings but also instrumental observations and captures of known

Manuscript submitted on February 12, 1980.

RNV. LYNCH III

luminous organisms. These reports concentrate on the spatial distribution of bioluminescence, andattempt to look at seasonal occurrences as well. Lynch (181 reviewed this literature and also discussedvertical distribution and diet variation of bioluminescence.

To be seen by the human eye, displays must be relatively bright. However, two instruments capa-ble of detecting light intensities well below the threshold of the human eye have been used in marinebioluminescence research. Bathyphotometers are designed to measure light beneath the surface of theocean. They may examine large or small volumes of water, may or may not provide their own sourceof stimulation, and may or may not be towable. Efforts are underway to design a floating bathyphotom-*2 eter, capable of operating remotely in the manner of current meters. The second instrument is the lowlight level image intensifier (LILIL). These instruments can be used on surface ships or airplanes, and

* satellite-borne instruments are even being contemplated. They remotely sense dim bioluminescence,enhance it, and provide a picture of the display on a television screen. A ship apparently dark to theeye may be readily identifiable with a LLLII. These two sensors are more fully discussed by Lynch [1]and Hickman, et a/. [71.

METHODS

For this report the studies on spatial and temporal distribution of bioluminescence cited abovewere reviewed. Reliance was placed on the statistical work of Turner 115,161 though he cited certaindeficientcies in his own reports. Since these works are compilations of individual reports ranging innumber from hundreds to thousands, it was not felt necessary to refer to the original individual reports.Tarasov (191 was consulted for Russian work before 1956 in the Black Sea. However, his analysis ofworld-wide distribution was ignored because of lack of accuracy in analyzing the data of Smith [8,91 andGlahn 1131 and because of lack of documentation of original Russian data cited (see comments in 1151).For information on the luminous displays in the Arabian Sea, Kalle 120,211, Tett and Kelly [13, andKelly and Tett 151 were consulted in addition to the works cited above. An analysis of the numbei andlocation of scientific studies of oceanic bioluminescence in Hickman, et aL (71 was also useful.

In estimating the frequency of bioluminescence encounters the author compared the number ofreports from a given area with a rough estimate of the shipping density in that area. Because of lack ofknowledge of exact shipping densities, a numerical analysis was not attempted. Because of the con-sensus that the Arabian Sea was the most luminescent region of the oceans, the author chose that ashis standard for hvery high frequency." Where shipping density was high, as in the Arabian Sea, but thenum~ber of reports was lower by approximately half, the author took this to be "high frequency."IAnother halving in the number of reports in dense shipping areas was taken to be "moderate fre-quency." Areas of dense shipping but few reports were considered to be "low frequency" areas.Overall, and especially in areas of little shipping, the author relied on his personal experience at sea,where he found bioluminescence in every location except one where measurements were attempted.Thus, the maps presented are somewhat subjective. It is unfortunate that existing data are too scantyto allow for a more reliable statistical presentation. However, the maps presented are of value inestimating the chances for encountering bioluminescence in a given area at a given time, provided thattheir limitations are understood.

RESULTS

Resuits are presented as a series of maps covering the Atlantic, Pacific, and Indian Oceans, withmore detailed maps for the Mediterranean Sea and Gulf of Mexico-Caribbean Sea. Four maps, one foreach season, are provided for each region. A fifth map is given to show the general density of reportsfor the region.

2

000 n4.

- - ."T .

NRL REPORT 8475

L iDISCUSSION AND CONCLUSIONS

Atlantic Ocean

Smith [8,9) analyzed the seasonal occurrence of bioluminescence in 10°-Marsden squares 145-151, which span the Atlantic from Nova Scotia to the coast of France. He found low levels ofbioluminescence •n winter throughout, and a tendency to peak in spring off Newfoundland and in sum-mer near the European coast. In the mid-Atlantic no seasonal peak could be distinguished. Glahn [131studied 10°-squares between the latitudes 0° and 60° and found a tendency for bioluminescence to peakin the spring and early summer in the high latitudes of the North Atlantic. Nearer the equator andthroughout the South, Atlantic occurrence appeared fairly even throughout the year. Staples [171 agreedwith Glahn. In addition, he observed that luminescence in the coastal waters of the British Isles andthe Scandinavian Peninsula was associated with dinoflagellate blooms, which tend to occur in late springand early fall. Lynch [11, while agreeing with the works cited above, noted that luminescence was oftenassociated with red tides off the coasts of New England and Florida. Red tides, which are nothing morethan unusually heavy dinoflagellate blooms, may occur throughout the year but are most frequent inlate spring and early fall. Near the equator all the above authors and Turner [15,161 agreed thatbioluminescence occurred fairly evenly throughout the year and at relatively high levels. Data for theSouth Atlantic are relatively sparse, but Smith, Glahn, Staples, and Turner all agreed that the area nearthe mouth of the Rio de la Plata was very bright throughout the year. Lynch has noted moderateluminescence off Namibia during three quarters of the year. In the Atlantic, no correlation has beendefinitely established between the occurrence of bioluminescence and any physical or meteorologicalphenomena other than dinoflagellate blooms. These analyses are reflected in Figs. I through 5.

Shipping in the North Atlantic between Europe and Nortn America, near the equator betweenSouth America and Africa, and along the continental coasts is very dense. Thus, considerable reliancecan be placed on the accuracy of the maps in these regions. In the far North Atlantic and most of theSouth Atlantic, shipping is much less dense. Therefore the maps are considerably less rellable in theseareas.

Gulf of Mexico and Caribbean Sea

Bioluminescence freqnency in the Gulf of Mexico is largely unknown, except along the coasts ofthe United States where it is frequently associated with red tides. Staples 1171 and Lynch [11 have indi-cated that bioluminescence occurred throughout the year, but less in winter than at other times. Smith[8,9] and Turner [15,161 counted relatively few reports for the region. 'Shipping was infrequent in thewestern Gulf, but high in the eastern Gulf and along the coast.

In the Caribbean Sea, shipping was dense throughout. Smith, Staples and Turner rountedmoderate numbers of reports in this region, although Lynch indicated high numbers. Staples andLynch indicated that bioluminescence occurred fairly evenly throughout the year. Three areas in theCaribbean showed constant high levels of bioluminescence. These were the shallow luminous bays ofPuerto Rico and Jamaica. In these bays permanent colonies of luminous dinoflagellates exist in bloomconditions. Data for these regions are shown in Figs. 6 through 10.

Mediterranean and Black Seas

In the Mediterranean Sea bioluminescence seemed to occur with moderate frequency throughoutthe year, according to Smith 18,9], Turner [15,161, and Lynch 111. Glahn [13] suggested that abioluminescence maximum occurred in spring, while Staples [171 indicated that a minimum occurred insummer. All the above authors agreed, however, that the western Mediterranean was brighter than theeastern, and that the area near the Strait of Gibraltar was especially bright.

3

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RV. LYNCH III

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Fig. I -Frequency of bioluminescence encounters in the Atlantic Ocean, Jan. through Mar.

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NRL REPORT 8475

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Fi. 6 - Frtequw o bkk rmh encomufte In the Out of Mexicoand Caribbean Sea, apr through June

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Fig.? 7-- FreQuenc of bioluminescahc encount in tw Gulf of Mexico

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5

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NRL REPORT 8475

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33

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250 1 . 3i5W 0 150E 30455Fig. 12 - Frequency of bioluminescnce encounters in the Mediterranean

and Black Seas, Apr. through June

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* RAE FRQUCNY 30

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15W 0 150E 300 45Fig. 13 - Frequency of bioluminescence encounters in the Mediterranean

and Black Seas, July through Sept.

I OW 0 15*E 300 *0

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Fig. 14 - Frequency of bioluminescence encounters in the Mediterraneanand Black Seas, Oct. through Dec.

12

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I HIGH FREQUENCY2 MOOERATE FREQL.NCY

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Fig. 15 - Frequency of bioluminescence reports in the Mediterranean and Black Seas

Pacific Ocean

Data for the eastern Pacific region are shown in Figs. 16 through 20; for the central Pacific inFigs. 21 through 25; and for the western Pacific in Figs. 26 through 30. Shipping in the Pacific is fairlydense along the continental coasts, around Hawaii and in Japan, the Philippines and Indonesia.However, there are vast areas, especiaijy scvutheast of New Zealand and the Society Islands, where ship-ping is extrenrely sparse. Likewise, reports of bioluminescence in the Pacific are relatively few, mostcoming from near the Panama Canal, along the coast of the contiguous United States, near Hawaii,Japtn and New Guinea, and in the South China Sea. This information is shown in Figs. 20, 25 and 30.

Glahn [131 performed a seasonal analysis on his Pacific data for comparison to his Atlantic dataand found bioluminescence minimal in both the North and South Pacific in their respective winters.These minima were ,iore pronounced at higher latitudes. Near the equator bioluminescence occurredfairly steadily throughout the year. These findings were supported by Staples [171 and Lynch [1], whoalso noted bioluminescence maxima along the California coast and around Japan in late spring and sum-mer. In addition, Lynh noted the frequent association of luminescence with red tides in both loca-tions. In the Sea of Okhotsk, Stukalin [101 observed bioluminescence all year around, even in brokenice fields in winter. However, there was a maximum in August through September.

Turner [15,161 and Staples [171 agreed that bioluminescence was fairly common in the SouthChina Sea. In addition, Turner performed a seasonal analysis for four regions within the South ChinaSea and for the sea as a whole. He found relatively constant levels of bioluminescence throughout theyear, with a slight apparent maximum in October through November. However, because of the smallnumber of reports available he considered this maximum an artifact resulting from the inadequacy ofthe data.

Indian Ocean jThe consensus of all who have studied the world-wide distribution of bioluminescence is that the I

Arabian Sea and nearby waters are the brightest regions in the oceans. The Bay of Bengal and the Straitof Malacca are also unusually rich in luminescence. Furthermore, this brightness extends throughoutthe year. But Smith [8,91, Glahn [131, and Anonymous [111 showed a bioluminescence maximum inlate summer and early fall. This maximum appeared to be correlated with the time of greatest current

13

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Fig. 16 -Frequency of bioiumhmmusence encounters inthEastern Pacific Ocean, Jan. through Mar.

14

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Fig. 19 - Frequency of bloluminewcence encounters inthe Eastern Pacific Ocan, Oct. through Dec.

17

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Fig. 21 - Frequency of bloluminescence encounten Iinthe Central Pacific Ocean, Jan. through Mar.

19 ý

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Fig. 24 - Frequenc of biobakmiswem encou s ntnhthe Ceftrl PckOmen. Oct. ftroamh Dec-

22

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Fig. 27 - Frequency of biolwumnescence encounters in the Western Pacificand Indian Ocean, Apr. through June

25

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Fig. 28 -Frequency of bioluminescence encounters in fte Western Pacificand Indian Ocean, July ftroubh Sept.

26

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28 ~ IC

17IHFROCC

NRL REPORT 5475

strength and upwelling due to the Southwest Monsoon, Staples (171, Tett and Kelly 131, and Lynch [1)suggested a secondary bioluminescence maximum in January through February, corresponding toupwelling induced by the Northeast Monsoon. However, both this upwelling and the consequent

bioluminescence are much less than that due to the Southwest Monsoon.

For the remainder of the Indian Ocean few reports are available. Lynch [I] analyzed the taxo-nomic data of the International Indian Ocean Expedition apd found bioluminescent organisms to bevery common throughout the studied region and at all times of the year.

A special word is in order About luminous displays in the Arabian Sea and its environs. This areais one of t~he richest in the world for displayr, of the 'phosphorescent wheel" and "milky seae types.Only the Strait of Malacca-South China Sea area approaches the Arabian Sea in terms of number ofdisplays. Neither the causative organism(s) nor the triggering mecharism i(s) of these displays havebeen identified. Herring 141 described several types of displays and discassd possible causative organ-isms. Turner 115,161 and Staples 1171 also described various types of displays, and Turner showed thatmost milky seas in the northern Arabian Sea occur in August, while Staples found that most phos-phorescent wheels in the same area appeared from April through June.

Many desctiptions of phosphorescent wheels exist, among them Tydeman [221 Termutelen [231,Rodewald [241, Kalle (20,211, and Hilder [251, who also offer tentative explanations for theiroccurrence. Tydeman [221 and Termndtelen [231 suggest that the wheels are due to optical illusionscreated by the motion of the ship and the interaction of the bow wave of the ship with existing wavesin a patch of bioluminescence. This theory has been mathematically explored by Verploegh [261, whoalso considered parallax resulting from the position of the viewer. Kalle 120,211 suggested that wheelsand "erupting balls are triggered by seismic disturbances from underwater earthquakes or volcanos.Hilder [271 observed bioluminescent streaks apparently triggered by ship radar. Because of thisobservation he suggested [251 that wheels are triggered by electromagnetic anomalies due to sunspotsor movement of ships through the earth's magnetic field. Rodewald (241 described an extremelyunusual phenomenon, phosphorescent wheels appearing above the sea surface. He suggested that thesewere caused by luminous microorganisms suckAd into the atmosphere with water droplets as aerosols.None of these suggestions has been proved. A discussion of wheels along with suggested research intotheir causes appears in Lynch [281.

STATISTICAL CONSIDERATIONS

All studies of the patterns of bioluminescence in the oceans have suffered from a bias towardheavily travelled sea lanes. A large number of reports from an area may represent a high level ofbioluminescence or it may represent the presence of many ships. Turner (15,161 made the first attemptto correct for this factor by defining a coefficient equal to the percentage of the world total of reportswhich came from a given area divided by the percentage of the world total of observing ships in thatarea. He derived his estimate of these percentages by limititl his data to that provided by the Volun-tary Observing Fleet from 1920-1938. These ships systematically made four sets of meteorologicalobservations per day and reported the data to the British Meteorological Office along with observationsof any unusual phenomena such as bioluminescence. By restricting his data base in this manner,Turner could know the exact numbers and locations of reporting ships. Thus, he could calculate hiscoefficient and eliminate the effects of ship traffic from his comparisons of the relative frequencies ofbioluminescence occurrences in given areas. Because of the limited number of reports, Turner felt thatthe smallest reliable unit for this kind of analysis was a 10°-Marsden square. Moreover, as Turnerpointed out, there are several defects in this method. First, it is based on the assumption that if twiceas many ships cross an area, they will send in on the average twice as many reports. Second, there isno way to correct for the passage of a ship through a comer of a square between observations or duringthe day. However, since this objection applies equally throughout the world, Turner felt that it did notinvalidate the method for comparing regions. Third, in places where ship traffic is very heavy, potential

29

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RN. LYNCHl IlI

observers may be fully occupied with ship handling and observations may not be made or recorded.2 Fourth, the correlation between the number of ships and the number of reports becomes unreliable

when the shipping density is low, because unavoidable inaccuracies in the shipping estimates becomesignificant at low densities.

Calculation of the coefficient is possible only if the data base is restricted to a fixed number ofships which make systematic observations. Acceptance of any reported observations makes impossible ~a reliable estimate of the percentage of the world total of observing ships in an area, because no reportwill be made if a ship observes nothing. Furthermore, observations are generally recorded and reportedfrom general shipping only if they are unusual and if the observer has the interest to report and knows

* where to report. The author personally knows of many observations which were not reported becausethe observers did not know where to go or because they had observed dim bioluminescence so often

* that they considered it routine. Thus, Turner's method is not suitable for analysis of reports from gen-eral merchant or military shipping because the irregularity of the reports creates built-in statisticalunreliability.

Now that instruments are capable of detecting bioluminescence not visible to the human eye,another source of statistical unreliability has been introduced. Observers using such instruments willobviously see more bioluminescence than those not using them. Thus, it becomes impossible to com-pare results between the two kinds of observers.

Tt-is report suffers from all of the above statistical problems. For that reason no attempt has beenmade to attach numbers to any of the areas of occurrence. outlined on the maps. A high frequency ofoccurrence simply means that one will likely encounter bioluminescence much more often thanmoderate frequency would suggest. However, to give some absolute value to the frequency categoriesused in the maps, the author suggests that the category of 'Moderate Frequency" means the constantpresence at night of instrumentally measurable bioluminescence, and that 'Very High Frequency* mightmean optically visible displays occurring once or twice a week. One should also realize that the bordersof the luminous regions categorized cn the maps are imprecise, and temporal changes occur graduallyand not abruptly on the first day of each quarter. The maps showing the frequency of reports havebeen included to give some indication of the reliability of the frequency of encounter data. Data fromareas with high reporting frequencies are more reliable than data from areas with low reporting frequen-cies. Thus, the encounter estimates from heavily travelled shipping lanes are good, in spite of the sta-

tistical problems, while those from little-xplored regions may be poor.

CORRELATIONS ICorrelations have been postulated between bioluminescence occurrences and regions of high

marine productivity 15,181. This is believed to be largely due to luminous dinoflagellates and occurs

most often in shallow waters. The connection between coastal displays off California and luminousdinoflagellates was shown early by Allen (191. Luminous dinofiaellates also occur frequently, but notI '*always, in red tides. However, the correlation with highly productive regions is not absolute. Kelly andTett [5i and Lynch (181 have cited exceptions to the theory. The relationship between productivity andbioluminescence may be highly complex [31, but its exact nature is currently unknown.

ACKNOWLEDGMENTS

I wish to thank John C. Moon, the artist who prepared the maps that comprise the bulk of thisreport. I also wish to thank the Navy Science Assistance Program which funded this report.

30

NRL REPORT 8475

REFERENCES

1. R.V. Lynch, "The Occurrence and Distribution or Surface Bioluminescence in the Oceans During1966 through 1977," NRL Report 8210, 1978.

2. E.N. Harvey, Bioluminescence, Academic Press, N.Y., 1952.

3. P.B. Teti and M.G. Kelly, "Marine Bioluminescence," oceanoS. Mar. Biol. Ann. Rev. 11:89-173,1973.

4. P.J. Herring, "Observations of Bioluminescence at Sea," Mar. Obs. 46:176-183, 1976.

5. M.G. Kelly and P.B. Tett, "Bioluminescence in the Ocean," in Bioluminescence in Action, P.J. Her-ring, ed., Academic Press, N.Y., pp. 399-417, 1978.

'. G.D. Hickman and R.F. Staples, "Bioluminescence of the World's Oceans: Annotated Bibliogra-phy," Applied Science Technology Report AST-R-03879, 1979.

7. G.D. Hickman, R.F. Staples, and R.V. Lynch, "Bioluminescence of the World's Oceans: TechnicalAsscL.sment," Applied Science Technology Report AST-R-080880, 1980.

8. H.T. Smith, "Phosphorescence of the Sea," Mar. Obs. 3:193-1%, 1926.

9. 1. Smith, "Phosphorescence of the Sea," Mar. Obs. 8:230-234, 1931.

10. " I.V. Stukalin, "Bioluminescence of the Okhotsk Sea," Vest. dal'nevost. Fil. Akad. Nauk SSSR';o. 9, pp. 137-139 (USHO Trans. 160), 1934.

11. .onymous, "Meerleuchten im Arabischen Meer," Seewart 8:12-18, 1939.

12. R.V. Lynch, "Bioluminescence in and near the Arabian Sea," NRL Letter Report 4351-78, 1980.

13. W. ('ahn, "Mcerleuchten im Atlantischen Ozean," Seewart 12:17-25, 1943.

14. V Murina, "Luminescence in the North Sea," Priroda, No. 12, p. 116 (USHO Trans. 22), 1954.

15. R.J. Turner, "Notes on the Nature and Occurrence of Marine Bioluminescence Phenomena,"National Institute of Oceanography Internal Report B4, 1965.

16. R.J. Turner, "Marine Bioluminescence," Mar. Obs. 36:20-29, 1966.

17. R.F. Staples, "The Distribution and Characteristics of Surface Bioluminescence in the Oceans,"Naval Oreaographic Office Report TR-184, 1966.

18. R.V. Lynch, "The Distribution of Luminous Marine Organisms," Proc. Naval Oceans Systems1Center Workshop on Bioluminescence (In press).

19. N.I. Tarasov, Luminescence of the Sea, Akad. Nauk SSSR, Moscow (Trans. M. Slessers, 1957, USNavy Hydrographic Office) 1956.

20. K. Kalle, "Die ritselhafte und 'unheimliche' Naturerscheinung des 'explodierenden' und des'rotierenden' Meeresleuchtens-eine Folge lokaler Seebeben?" Deutsche Hydrog. Zelts. 13:49-77,1960.

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21. K. Kalle, "Wheels of Light," Sea Frontiers 1S(2):116-122, 1969.

22. G.F. Tydeman, "Phosphorescent Wheels," Mar. Obs. 9:32, 1932.

23. J.W. TermUtelen, "Phosphorescent Wheels," Mar. Obs. 20:229.232, 1950.

24. M. Rodewald, "Phosphorescent Wheels," Mar. Obs. 24:233-234, 1954.

25. B. Hilder, *Marine Phosphorescence and Magnetism," Navigation (J. Aust. Inst. Navigation)1:43-60, 1962.

26. G. Verploegh, "The Phosphorescent Wheel," Deutsche Hydrog. Zeits. 21:153-162, 1968.

27. B. Hilder, "Radar and Phosphorescence at Sea," Nature, London 176:174-175, 1955.

28. R.V. Lynch, "Problems and Opportunities Connected with Marine Biological Phenomena," inAdvanced Concepis in Ocean Measurements for Marine Biology. F.P. Diemer, SJ. Vemberg, and -D.Z. Mirke., eds., University of South Carolina Press, Columbia, SC, pp. 245-255, 1980.

29. W.E. Allen, "'Phosphorescence' in the Sea," Naut. Gaz., Aug. 26, pp. 10 and 16, 1939.

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