TR-184

TECHNICAL REPORT

THE DISTRIBUTION AND CHARACTERISTICS
OF SURFACE BIOLUMINESCENCE
IN THE OCEANS

ROBERT F. STAPLES
Biological Section, Environment Branch

Oceanographic Analysis Division

MARCH 1966

/
/ TH3 U.S. NAVAL OCEANOGRAPHIC OFFICE
; ; WASHINGTON, D.C. 20390
pw TR [64
Price 90 cents

MBL/WHOI

AOA

FOREWORD

Bioluminescence of the sea generally has been regarded as a
mysterious phenomenon, and observations in the past for the most
part have been fragmentary and unreliable. It is only in recent
times that systematic observations and explanations of the phenom-

enon have become available.

In an effort to summarize the data already in existence, the
U. S. Naval Oceanographic Office has undertaken an intensive
literature search which included reports extending back almost a
century. As expected, reports of displays for the most part came
from shipping lanes and included very little information from less

frequented areas.
This study is published as part of the Oceanographic Office

program in marine biology. It is intended as a small contribution
to a field that has been sadly neglected for a long time.

tut Pritiie

Auted) Dstt D. WATERS, JR.
Rear Admiral, U. S. Navy

Commander
U. S.- Naval Oceanographic Office

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CONTENTS

INoprenoisal G Sagara "S S'S Mas MOMS aia Selon ana rere
IGRS Gir Joanna lg 6 GO SoM etd Go GBs 696 Gc
bieireriigenl REGIMES 6 56 6 0 oO Oooo Oo
Luminescent Marine Organisms . RS rere Sans i
Types of Bioluminescent Tagyeilare SG Ot Re Game
Observation Reports ...... « oe eh haatey ss
North Atlantic Ocean and Adjacent Gone oom one
Southern North Atlantic and South Aeaeunete Oceans

Northern Pacific Ocean and Adjacent Seas .....
East-Central and South Pacific Ocean .......
Western South Pacific and Eastern Indian Oceans .
Central and Western Indian Ocean and Adjacent Seas
"Phosphorescent Wheel" and "Wave" Displays ... .
Rioillaye ResHIOME 6 o 6 6 6 6 6 0 oOo OO OOO
BLIGE, 6 o 6 6 oO 6 OOO OO OO OO

Figure

LIST OF FIGURES

Seasonal Distribution of Bioluminescent Displays,
Northern Atlantic Ocean and Adjacent Seas 5

Seasonal Distribution of Bioluminescent Displays,
Southern North Atlantic and South Atlantic Oceans
(Gamera Toe Mw) 6 B do 6 6 eyowe ib Sc

Seasonal Distribution of Bioluminescent Displays,
Southern North Atlantic and South Atlantic Oceans
(ina throusheiecember)) vey remem ier nee

Seasonal Distribution of Bioluminescent Displays,
North Pacific Ocean and Adjacent Seas 2 3-0. =

Seasonal Distribution of Bioluminescent Displays,
East-Central and South Pacific Ocean .....

Seasonal Distribution of Bioluminescent Displays,
Western South Pacific and Eastern Indian Oceans

Seasonal Distribution of Bioluminescent Displays,

Central and Western Indian Ocean and Adjacent Seas

Distribution of "Phosphorescent Wheel" Displays .. .

vi

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35
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THE DISTRIBUTION AND CHARACTERISTICS
OF SURFACE BIOLUMINESCENCE IN THE OCEANS

HISTORICAL RESUME

A fully developed luminescent sea is one of the most striking natural
phenomena that mariners or sea travelers can witness. Many reports in the
form of log entries or as part of various written narratives in travel
books or cruise reports have referred to luminescent seas since ancient
times. During past centuries these strong displays of light often were
viewed as having mystical meanings and were identified with the super-
natural. In the South Seas, for example, islanders considered the fire
they used on land as originating from the brilliant luminescent seas
they often encountered during their travels among the numerous islands.
Whatever ancient peoples thought of bioluminescence, they left little
record of their observances, except for a few poetic references and some
more disciplined records by Aristotle.

Descriptive reports of bioluminescent seas of a more scientific nature
increased after the 15th century. Various sea captains, whether on ex-
ploring, military, or commercial ventures, made numerous records of lumin-
ous seas from almost all the oceans of the world. One of the earliest
descriptive reports on file concerned a "burning and glittering light of
the sea ... as though all the Sea ourer had beene burning flames of fire."
This luminous condition of the water was observed by the explorer John
Davis in the 17th century near Ascension Island. The terms "burning sea"
and "milky sea" often appeared in the records. The ocean surface, when
strongly lighted by the phenomenon, appeared to be burning, to give off
_ flames as if on fire, and to contain what appeared to be "smoldering
coals." Sailors to this day have called such manifestations "phosphor-
escent seas.” We now know this term to be a misnomer; the luminescence
of the sea is bioluminescence, or biological light caused by living
organisms, rather than phosphorescence, which in the strict sense results
only from the irradiation of inanimate substances by radiant energy.

"Milky seas" were observed in various oceans, mostly tropical, and
were particularly intensive in the Indian Ocean, Arabian Sea, and the
seas of the Indonesian Archipelago. Stavorinus (1798), a Dutch navigator
on a voyage to the East Indies, described this phenomenon near the coast
of southern Saudi Arabia.

"On the 30th (January) we met at night a very singular
appearance in the color of the sea. It assumed so great

a degree of whiteness that it was perfectly like milk...

as the evening twilight diminished, it became whiter and
increased gradually in whiteness till nine o'clock when it
was so white that the whole sea appeared as if covered with
a white sheet, or exactly like the appearance in the night
time of a flat country overspread with snow.”

This resemblance to a snow-covered plain led the Dutch to call such
appearances the "winter ocean" or "the white water," the latter term
particularly suited in describing the snow-white luminescence often ob-
served during late winter in the Banda Sea.

Systematic collections of reports concerning bioluminescent seas
were not made until comparatively recent times. During the latter half
of the 19th century, articles published by the Meteorological Offices of
both England and Germany included records of bioluminescent displays,
primarily in the Atlantic Ocean. More recently the incidence of bio-
luminescence in the Atlantic was charted by Smith (1931). His charts
were based upon numerous observations of bioluminescence reported in the
British publication "The Marine Observer." The seasonal distribution of
displays in the Arabian Sea and adjacent regions was reported in the
publication "Der Seewart" (1939). Turner (1965) has recently written a
comprehensive report on the nature and occurrence of marine biolumines-
cence. This report was received as our publication was going to press.

LUMINESCENT MARINE ORGANISMS

Plankton organisms chiefly are responsible for bioluminescence in
the sea. The smallest forms are luminescent bacteria which usually live
on decaying matter or within various marine animals. However, with a
supply of the proper nutrients, luminescent bacteria can develop in great
masses in the sea, causing a general bluish green glow in the water. The
glow is usually diffuse and barely detectable, although exceptionally
bright displays caused by luminous bacteria occasionally are observed in
coastal regions near the outflow of large rivers. The light given off
frequently outlines the current front where the river and ocean meet.
Soviet investigators believe that luminescent bacteria have a much larger
role in producing bioluminescence than that considered by American or
European scientists.

Most bioluminescence in the oceans is produced by one-celled organisms
called dinoflagellates. Of these small luminescent organisms, members of
the genus Noctiluca are associated most frequently with strong displays of
light. Although individuals are just visible to the eye, they can develop
in such prodigious numbers that they color the water pink or red by day.
Noctiluca is particularly abundant in coastal waters and at night imparts

a rather brilliant greenish glow to the water when mechanically disturbed.

tH

Fishermen in some regions have long used the cry of "Noctiluca!" to bring
fiery illuminations to the attention of their fellow fishermen, regard-
less of the organism causing the luminescence. Holder (1887) observed a
display caused by vast accumulations of Noctiluca. He reported:

"On the 10th of April, in the evening, the sea in the roadstead
of Simonstown, Cape of Good Hope, presented an extraordinary
phosphorescence of a most vivid character. At whatever points
the phosphorescence was greatest, the water was colored on the
surface as red blood and it contained such an immense quantity
of little globules (Noctiluca) that it had the consistency of
syrup, the globules consisting of more than half the volume of
the seawater."

Other dinoflagellates also impart luminescence to the ocean.
Pyrocystis, often confused with Noctiluca, is responsible for many exten-
sive displays on the high seas. Organisms such as Ceratium, Peridinium,
and Gonyaulax cause rather bright surface luminescence. Gonyaulax also
is one of the prime causes of "red tide."

Other organisms responsible for luminous waters include crustaceans,
such as ostracods, copepods, and euphausiids. Most of the displays
caused by crustaceans are seen in colder waters and only rarely in tropi-
cal waters. The light emitted appears to twinkle at a distance because
of abrupt flashing by each individual and is usually blue or green.

Luminescent jellyfishes (Medusae) also cause many displays. Large
shining round or oval spots of light may appear in the water and the
resulting luminous sea can be very bright and cover a large area. One
of the most spectacular forms is the large luminescent medusa Pelagia
noctiluca. When touched lightly the whole surface of the organism
starts to luminesce, first at the point of contact, then spreading out
to the umbrella and tentacles.

Ctenophores (comb jellies) are almost all luminescent, giving off
a greenish glow. Luminescent transparent tunicates such as Salpa or
Pyrosoma are responsible for some of the most spectacular displays re-
ported by mariners. The latter organism is abundant in warm waters, and
the light produced by large concentrations of colonies has been described
by Thomson (1877) while a member of the CHALLENGER Expedition.

"After leaving the Cape Verde Islands and going south in
August, between 14° and 22°w and 5° and 17-N, the sea had
been every night a perfect blaze of phosphorescence, the un-
broken part of the surface appeared pitch black, but wherever
there was the least ripple the whole line broke into a
brilliant crest of clear white light. Near the ship the black
interspaces predominated, but as the distance increased the
glittering ridges looked closer until towards the horizon as
far as the eye could reach, they seemed to run together and
to melt into one continuous sea of light. The wake of the
ship was an avenue of intense brightness. It was easy to
read the smallest print sitting at the after-port in my cabin;
the bows shed on either side rapidly widening wedges of radi-
ance, so vivid as to throw the sails and rigging into distinct
lights and shadows. The first night after leaving San Iago,
the phosphorescence seemed to be chiefly due to large Pyrosoma
of which we took many specimens with a tow net, and which
glowed in the water with a white light like that from molten
iron.”

The slightest touch at one end of a colony of Pyrosoma can cause blue
light to advance along the structure, each individual organism lighting
in turn, until the whole luminesces brilliantly. Two popular names for
this organism are the "fire body" and "fire cylinder." Various colors of

Pyrosoma luminescence have been reported, such as red, orange, yellow,
and white light, but these result when the organisms are overexcited or
dying. The light normally given off in the sea is bluish green or green.
In colder water luminescent salps often are in great abundance and may
be present as individuals or in great chainlike aggregations. They
luminesce blué or green.

Except for the above organisms, the majority of luminescent forms in
the ocean rarely occur in great enough numbers to create marked surface
displays. However, some of these may create displays at certain times
and places. Some seaworms may luminesce during their spawning periods
when they swarm in surface waters, and the resulting light may be very
strong. The lights Columbus observed on the night preceding his landing
on San Salvador may have been caused by the luminescent worm Odontosyllis.
Other forms, such as the deep-sea squid Watasenia scintillans (Japanese
firefly squid), may congregate in large numbers on the surface during the
spring, causing displays. Microscopic radiolarians occasionally may im-
part a weak luminescence to the ocean.

TYPES OF BIOLUMINESCENT DISPLAYS

Bioluminescent displays may be classified according to their appear-
ance. The observations presented in this report have been grouped into
three general descriptive categories based, with some modifications, on
the types proposed by Tarasov (1956). We have named these three: sheet
type, spark type, and glowing-ball or globe type.

The sheet-type display is the most common type observed in coastal
waters and is caused by masses of dinoflagellates or bacteria. Tarasov
(1956) referred to this luminescence as spilled or "milky" bioluminescence.
The water may appear dully luminescent, and the individual points of light
are not resolvable; that is, they give a sort of luminous cloudlike
appearance in the water. The color usually is green or blue and in many
displays appears white when the organisms are present in great numbers.
From this latter appearance the term "milky sea" has been derived. Dense
and extensive concentrations of large organisms, such as Pyrosoma, lumi-
nescing at the same time, may appear as a sheet-type display, instead
of the flashing appearance noted when they are less concentrated. The
same effect is produced by large concentrations of euphausiids, as
Murray (Thomson and Murray, 1885) reported in August 1880 while in the
Faeroe Straits (Faeroe-Shetland Channel). He noted: "large spots and
long bands of milk-white water.” The luminescence was caused by count-
less numbers of Nyctiphanes norvegica, a luminescent euphausiid.

Spark-type displays are created by large numbers of luminous
euphausiids or copepods. This kind of luminescence occurs most often
in colder waters and only when the waters are disturbed. Luminescing
of the organisms gives the ocean surface a scintillating appearance.
The luminescence produced usually is brilliant blue or white, and the
light flashes are just resolvable to observers from the deck of a ship.

Glowing-ball- or globe-type displays are seen most frequently in the
warmer waters of the world. The ocean may appear as if full of balls or
discs of light, some flashing brightly as they are disturbed and others
dimming after the initial stimulus has ceased. The flashes or pulsations
of light may range in size from a few centimeters to a number of meters
in diameter, depending upon the size of the organisms. The light given
off usually is blue or green; occasionally, displays of white, yellow,
orange, or red have been reported. The light rarely is continuous and
may be noted from afar by its flashing appearance.

Combinations of either two or all three types of displays often occur.
The light given off may consist of flashing luminescence against a cloud-
like luminescence in the water or a mixture of glowing-ball- and sheet-
type luminescence. In some observations, especially in higher latitudes,
the scintillations of luminous crustaceans may be seen as they graze on
the edges of large concentrations of luminous dinoflagellates, that is,

a spark-type display fringing on a sheet-type display.

Exotic light formations like "phosphorescent wheels," undulating waves
of light, and bubbles of light appear to be separate and distinct from
all three types discussed above. "Phosphorescent wheels" and "waves" of
light are discussed in another section.

OBSERVATION REPORTS

In preparing the charts used in this report the writer had access to
files in the Biological Section of the U. S. Naval Oceanographic Office
containing approximately 3,000 individual reports of displays throughout

most of the oceans of the world. These reports date primarily from the
beginning of the 19th century to the present time; however, a few reports
are many centuries old, and these have been included in the overall pres-
entation where they were considered useful.

Most of the reports of displays have been obtained from publications
such as the British "The Marine Observer,” the “Hydrographic Bulletin"
of the U. S. Navy Hydrographic Office, and the "Notices to Mariners"
published jointly by the U. S. Coast Guard and the U. S. Naval Oceano-
graphic Office. In addition to the above publications, numerous scientific
eruise reports, especially narrative logs, provided many reports. Reports
from books of ocean travels also were included. The works of Harvey (1952),
Nicol (1960), and Tarasov (1956) were of special value.

As with many types of oceanographic and hydrographic data, a bias
exists in the geographic distribution of bioluminescence observations,
inasmuch as the majority of reports are from frequently traveled shipping
lanes. Consequently, reports of displays were unavailable or few in
number for many regions. When the available reports are all plotted, they
may give an erroneous representation of the seasonal distribution of bio-
luminescence in many places. Therefore, additional information such as
published general descriptions of bioluminescence, ecological factors,
distribution of organisms causing bioluminescence, and many other types of
miscellaneous data were used to provide a more comprehensive coverage of

the phenomenon.

NORTH ATLANTIC OCEAN AND ADJACENT SEAS (FIGURE 1)
Coastal Waters of Eastern United States

Bright displays (mostly sheet- and glowing-ball types) have been
observed in coastal waters of the eastern United States during spring and
late summer. Euphausiids and copepods often appear in large concentrations
capable of causing strong spark-type displays in the Gulf of Maine during
the summer. Ctenophores, such as Pleurobrachia, the sea gooseberry, have
been observed in large numbers in the Gulf of Maine and along the coasts
of southern New England. Dahlgren (1915) reported some observations of a
friend from Harpswell, Maine concerning bioluminescent water. The water
luminesced because of large concentrations of ctenophores, which glowed
with a green light as they were struck or entrapped within a towed net.
Diffuse surface luminescence has been observed during the spring and late
summer months in Long Island Sound and Narragansett Bay.

Farther south in Barnegat Harbor, New Jersey occasional displays
have been observed. These often have been bright enough to attract the
attention of observers on shore, as in the autumn of 1963 when large
numbers of luminescent copepods and dinoflagellates were present in the
water. Dahlgren (1915), while traveling in Chesapeake Bay, described
the vivid "green fire" that occurred as his ship sailed through water
containing large numbers of luminescent dinoflagellates. The light
emitted was strong enough to come through the porthole and reflect from
the ceiling of a stateroom.

Gulf of Mexico

The Gulf of Mexico contains a tropical plankton, including many
bioluminescent forms. Displays are as intense as in more northern waters,
although not as frequent. Most of the bioluminescence reported has been
observed in coastal regions or over shallow depths as in the Straits of
Florida. Agassiz (1888) noted both Pyrosoma and Salpa in coastal waters.
The pyrosomes were not as common as in Indian Ocean waters, and the dis-
plays caused by these forms were much weaker. Salps, according to
Agassiz, were larger in size than in more northern waters; the light
given off by them was bright green, supplemented by a bluish light from
other smaller salps.

"Red tides" often are noted along the west coast of Florida and in
Texas coastal waters during the summer. Connell and Cross (1950) described
the appearance of a "red tide" containing large numbers of a particularly
virulent marine dinoflagellate, Gonyaulax. At night the water appeared
luminescent. This condition has been observed in June and August, usually
after rains along the coast. Noctiluca also has been observed in the
sulf during March. This organism may be present in great numbers along
the Mexican coast and over Campeche Bank in the spring.

Along the north coast of Cuba the ocean often is strongly illuminated.
Fishermen make use of this bioluminescence for tracking fish as the fish
swim through the water. In Havana Harbor the water often is bioluminescent
during the cool December and January nights.

LEGEND
© JAN., FEB., MAR.
® APR., MAY, JUNE
+ JULY, AUG. , SEPT.
* OCT., NOV., DEC.

EACH SYMBOL REPRE-
SENTS ONE DISPLAY
REPORT.

FIGURE 1 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, NORTHERN
ATLANTIC OCEAN AND ADJACENT SEAS
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Open North Atlantic

The percentages of total observations of bioluminescence from the
open ocean of the North Atlantic west of 30°W for three-month periods are
as follows:

January through March----- ahd
April through June-------- 38%
July through September---- 27%

October through December-- 11%

The percentages of total observations for the three types of displays in
this region are as follows:

Sheet type----------------
SSHNE TYING 2s SSS SSS sas 144%
Glowing-ball type--------- 34%

The most luminescence occurs in close proximity to regions in which
"mixing™ occurs, such as the front between the Gulf Stream and the cold
Labrador Current. Sheet-type bioluminescence predominates in the spring;
both sheet- and glowing-ball luminescence occur with equal frequency in
summer in the western North Atlantic.

Sheet-type displays, which are predominant in the region of "mixing,"
become less frequent eastward from 30 W, whereas glowing-ball displays
gradually increase in this direction. Glowing-ball displays have been
reported 54 percent of the time in this portion of the North Atlantic,
-most frequently in the spring. Sheet-type displays have been recorded
about 27 percent of the time and are most often seen from April through
September. Spark-type displays have been observed 19 percent of the time,
mostly from July through December.

The Sargasso Sea, noted for its floating vegetation, is, however, an
impoverished area with regard to other forms of marine life. As a con-
sequence, very little bioluminescence has been observed there. About
65 percent of the few reported observations were made from January through
June and 35 percent during the other half of the year. Although few in
number, some very strong displays have been reported, especially on the
fringes of this vast eddy. In 1905 observers on board the scientific
yacht PRINCESS ALICE observed a bright luminescent wake as they sailed
through the western portion of the Sargasso Sea. The luminescence was
globular in form and was caused by large numbers of Pelagia noctiluca, a
luminescent medusa noted for its green luminescence.

In the coastal waters of various island groups, such as the Azores
and the Canaries, luminescent displays occur during all seasons. Most
of the luminescence observed is caused by large shoals of jellyfishes
and tunicates carried into the area by prevailing currents. Around the
Canary Islands, bioluminescence has been observed most frequently during
the spring and autumn, whereas in the vicinity of the Azores the majority
of displays occur from April through December.

Atlantic Coastal Waters of NW Africa, Spain, Portugal, and France

Along the Atlantic coast of northwestern Africa and in the western
approaches to the Strait of Gibraltar, displays occur during the spring
and autumn months. On the Moroccan coast, in such places as the Bay of
Agadir, there also is an abundance of luminescent organisms during the
winter. In February 1960 these organisms luminesced brightly during the
earthquake which caused so much destruction on land. In October and
November 1955 moderately intense displays were noted near Casablanca,
Morocco. The light was bright enough to cause a "milky" glow which ex-
tended 2 feet from the sides of various oceanographic instruments as they
were raised and lowered into the water. These particular displays were
due to large concentrations of luminescent protozoans primarily, although
large flashes of light by ctenophores or jellyfish could be observed when-
ever an instrument jarred these organisms.

Some very spectacular displays occur in various bays of western Spain
and Portugal. Pinto (1949) gave an account concerning intense luminescence
and "red water" near Lourinha, Portugal in September 1944 caused by a bloom
of the dinoflagellate Gonyaulax. The’luminescence caused some panic among
local fishermen according to Pinto. The light given off was blue-green,
and the intensity increased when the water was disturbed. Displays such
as these may occur along the western Iberian coast from April through
December, although the most vivid displays are seen in the late summer
and early autumn. One of a number of wartime reports concerning bio-
luminescence and submarine tracking comes from this region. In November
1918 the last German U-boat (U-34) to be destroyed during World War I
was easily tracked because the water was so “phosphorescent” at the time
that the submarine could be seen moving under the water "glowing" and
outlined by "sea fire."

In the Bay of Biscay French and Spanish fishermen utilize summer
bioluminescence to track fish at night. The trails left and the peculiar
flowing movement of fish schools through luminescent waters provide them
with evidence as to the type of fish present. Noctiluca, so common to
most of the coastal regions of the world, causes bioluminescence in the
various bays and inlets along the coast of France. Quatrefages (1850),

a French naturalist, noted the intense luminescence of large concentra-
tions of this particular pinkish organism in water near such ports as
Dieppe, St. Malo, Bréhat, and Ostend (Belgium). These organisms appeared
in the surface waters from July through September.

English Channel and North Sea

In the English Channel, bioluminescence is most evident from April
through November; however, displays can occur during all seasons. Winter-
time bioluminescence is not unusual in this region, especially along the
English coast. For example, glowing forms of fishes and the long lumin-
escent trails they leave in the water have been observed in the dark
surface waters during February near Plymouth. "Brilliant green lines"

10

and "balls of phosphorescence" were noted by observers on board the SS
BEAVERGLEN in June 1957 off Beachy Head, England. Off Lands End, Cornwall
very bright light has been noted in the late summer, when shoals of
luminescent jellyfishes are brought in by currents.

Although many oceanographic and fisheries investigations have been
conducted in the North Sea, very little information is available con-
cerning bioluminescence in this region. As might be expected in
temperate regions such as this, the greatest peak of bioluminescent
activity appears to take place from April through June and a smaller
peak from late August through November. Bioluminescence generally is
restricted to shallow coastal regions or to the waters over the Dogger
Bank, although some strong displays can occur some distance from the
coasts over deeper regions.

The majority of the references are to spark-type bioluminescence.
Murina (1954) observed very intense bioluminescence east of the Orkney
Islands in August 1953. The surface water appeared to be full of
"sparkling emerald dots." Further investigation showed that a lumin-
escent copepod, Metridia lucens, was responsible for the light.

Various species of luminescent euphausiids also contribute to the spark-
type luminescence observed in late summer and early autumn. Euphausiids
may be present in large numbers in the numerous inlets and firths along
the Scottish coast, where they remain at depth during the day and migrate
to the surface layers at night, creating displays of some brilliance.

Sheet-type luminescence occurs in the late spring and late summer.
The light given off usually is produced by large numbers of luminous
-dinoflagellates, most prominently Ceratium, Peridinium, and Noctiluca.
The last organism causes bioluminescence in coastal regions in late
spring, and the other two organisms are responsible for late summer
displays. Noctiluca develops in such great numbers at times that it
constitutes up to a of the water volume.

Glowing-ball bioluminescence may be observed in such regions as the
Thames Estuary when influxes of luminous medusae occur. Farther to the
north great shoals of salps appear in late summer and early autumn at
the northern entrance to the North Sea and at times extend into the
central North Sea. Such large concentrations often create strong
glowing-ball displays.

Ocean Waters West and North of the British Isles

Although no specific reports are on file it is known that many
types of bioluminescent organisms cause displays in the ocean west of
the British Isles. Displays have been observed in coastal waters during
all seasons, although most of the bioluminescence appears during the
late spring, late summer, and early autumn. Scott (1920) noted that
although Noctiluca reached a peak of abundance in the coastal waters of
the Irish Sea in late summer and early autumn during mild winters as in

1919, this organism could be found in the water within Barrow Channel,
developing in great enough concentrations to discolor the water and to
eause strong sheet-type bioluminescence. One observer noted that during
the winter along the rocky coast of northwestern Scotland the sea was
luminescent. Every stroke of an oar produced a myriad of "green sparks”
in the disturbed water, and the wake luminesced for some distance
astern. Shoals of jellyfishes, salps, and ctenophores are responsible
for much of the summertime bioluminescence in the Irish Sea. Farther
north along the Scottish coast some displays due to dinoflagellates

and euphausiids often are observed in September. Swarms of herring
create bright luminous trails off the Hebrides Islands as they swim
through water containing luminescent copepods and dinoflagellates.

In the waters around the Faeroe and the Shetland Islands, displays
are most frequently observed from May through late October. The dis-
plays often are caused by euphausiids, which are common and at times
appear in dense concentrations throughout the region in spring, summer,
and autumn. The first aggregations of Salpa generally are sighted
inside the Faeroe-Shetland Channel in May and June. The water often
is so full of these organisms that it appears opaque, and the lumin-
escence given off at night can be very strong.

Norwegian and Greenland Seas

Strong displays can be expected during the year in the coastal
waters of Iceland and Norway and in regions where the warm North
Atlantic Current meets cold currents from the Arctic Ocean. The sea
ice also may be luminous because of entrapped luminous organisms.
Flashes of light have been seen when sea ice along the west coast of
Norway was broken or jarred. This luminescence continues in the melt
water. Copepods and dinoflagellates appear to be the organisms frozen,
but not dead, within the ice.

In the offshore waters of the Norwegian and Greenland Seas, Russian
observers have noted a considerable amount of bioluminescence in March
and April. In June and July this bioluminescence becomes stronger and
often extends over large regions. The organisms responsible probably
are dinoflagellates, most likely Ceratium or Peridinium. They can
create luminescence in ice-free waters along the coast of eastern
Greenland during the year. In August Metridia lucens is abundant in
Denmark Strait, and as a result displays of sparkling green light have
been observed in the various ice-free fiords along the Greenland coast.
Occasional displays also occur along the coast of Greenland and Iceland
from early summer through early autumn. Reports dating back to the
early 18th century make reference to the red discoloration of water in
southern Icelandic fiords, which turned into "sea fire" at night. Dis-
plays such as these are unusual for regions in such high latitudes.

Mediterranean Sea
The western basin is the most luminescent part of the Mediter-

ranean Sea except for the Adriatic Sea. Most of the displays observed

12

in the western Mediterranean occur from October through June. Sheet-
type displays predominate, with an occasional glowing-ball display.
Luminescent salps, pyrosomes, and medusae are responsible for many of
these displays. Pelagia noctiluca is especially plentiful in the sur-
face waters of the western Mediterranean. Salps often almost fill the
water; the organisms frequently form long chains and may emit long
lasting and rather intense light when disturbed. Noctiluca may be
present in great abundance, tinging the water red by day and luminescing
brightly by night.

During one of the many cruises of the Norwegian research vessel
MICHAEL SARS in April 1909, Murray (1912) noted that the sea surface
for several nautical miles in the Sea of Alboran was full of "phospho-
rescent" Noctiluca. The concentration was so great that the water was
like a "broth," and in the evening "the sea resembled a star-spangled
sky, and the wires following the vessel looked like gleaming stripes."
A more recent report from the SS EXANTHA in January 1951 bears out the
fact that strong sheet-type luminescence can be expected in this region.
The water was "phosphorescent" for several nautical miles along the
coast of southern Spain. The ship created a "luminous sheet" so in-
tense that it was possible to read by the light emitted. The cause
of this luminescence was not ascertained; however, it was of the type
produced by dinoflagellates such as Noctiluca. Very intense biolumin-
escence has been observed during the summer along the coast of Algeria.

Luminous euphausiids are responsible for many spark-type displays
along the coast of southern France and northwestern Italy. During the
winter months these organisms may appear in large swarms, coloring the
-water red. At night the light given off is sparkling and bluish white
in color, and when the organisms all light, the water takes on a "blue
radiance."

To the south, swarms of Noctiluca occur in the Bay of Naples in
August and September. Ehrenberg (1959), a German biologist, noted the
long luminescent wake created by these organisms as his boat traveled
through the water of the bay. He also noted that the fronds of a
common brown seaweed (Fucus) were covered by these tiny luminescent
forms, the fronds appearing to glitter when disturbed. Bioluminescence
occurs most frequently during the spring and summer months in the Bay
of Naples.

In the northern Adriatic Sea, bioluminescent displays are observed
more often than in any other part of the Mediterranesn. The lumines-
cence observed in the coastal waters here is most pronounced from April
through September. In April and May water near the mouths of the Po
may become very luminescent; the development of luminous organisms is
increased by the nutrients washed from land during the high springtime
runoff. At times bioluminescence may be produced by bacteria or proto-
zoa. Apparent bacterial luminescence usually is found in fiordlike
inlets along the northern Yugoslavian coast, where much decomposition
and stagnation occur.

13

In the Gulf of Trieste a luminescent slime often develops in the
late summer. The slime is yellow and ropy and adheres to objects in
the water. Therefore, it is not unusual to see pilings, rocks, and
bottoms of anchored boats luminesce yellow-green when covered by this
slime. The slime probably is produced by luminescent bacteria,
although dinoflagellates also have been suggested as the causative
organism.

Little information is available concerning bioluminescence in the
eastern Mediterranean. Two reports indicate wintertime bioluminescence
due to masses of copepods in the Aegean Sea. Some rather extensive
sheet-type bioluminescence occurs along the Egyptian, Israeli, and
Lebanese coasts in July, August, and September. Seaward from the Nile
Delta, fishermen locate fish by the luminescence stimulated by the fish
swimming at this time. Bioluminescence is most frequent during this
period because of the large development of luminescent plankton
triggered by the enormous quantities of nutrients carried seaward by
the flooding Nile.

Black Sea, Sea of Azov, and Caspian Sea

Tarasov (1956) states that luminescence can be observed through-
out the year in the Black Sea and that it reaches a maximum intensity
during the autumn months. Near Sevastopol Gonyaulax frequently causes
a red discoloration of the water. At night the waves are capped with
brilliant bioluminescence, and the surf gives off vivid light as it
breaks against the shore. In the Bay of Odessa very intense bio-
luminescence has been observed in the early summer, probably most
frequently near the mouths of rivers such as the Dnieper and Dniester.
Noctiluca is the dominant form in the summer plankton of the Black Sea.
While normally a coastal form, it occurs in large concentrations in the
open parts of the Black Sea. The light emitted by these protozoans is
augmented somewhat by large numbers of luminescent ctenophores during
the summer.

According to Zenkevitch (1963) the Sea of Azov is extremely pro-
ductive. The production reaches a maximum in the summer and early
autumn, and luminescent forms develop in such large concentrations
that they create sheet-type luminescence at this time. Sea worms also
develop in great numbers in the summer and create luminescence as they
move through concentrations of luminescent dinoflagellates.

Bacterial bioluminescence has been reported from the northwestern
part of the Caspian Sea. Zhirnov (1955) reported strong luminescence
near Astara in July 1952. The bioluminescence was evident in the
water for over a week and appeared to be due to tiny organisms visibly
luminescing even during the daytime. The organism causing the lumin-
escence was not determined.

Skagerrak, Kattegat, Belts, and Baltic Sea

In the Skagerrak considerable bioluminescence may be observed in
the early spring. Observers on board the MV LAKSA reported a display
off southern Norway in April 1963 which was determined to have been
caused by luminous dinoflagellates. Pleurobrachia, a luminescent
etenophore, is very abundant in the Skagerrak in May. Numbers of this
small transparent organism appear much like beads of green light when
viewed in the water. This ctenophore is especially prevalent inside
Oslo Fjord in the spring. Shoals of luminescent medusae may be en-
countered in the various fiords and inlets along the southwest coast
of Sweden during the summer. Sheet-type bioluminescence has been ob-
served in the various extensions of Oslo Fjord in August. Also, heavy
concentrations of luminous dinoflagellates are common in this fiord
during late summer. Euphausiids and copepods often create displays of
some intensity within the Skagerrak during late spring and summer.

Extensive displays of light are not common in the Kattegat, Oresund,
and the Belts. Some large concentrations of the tiny tunicate
Appendicularia are responsible for green luminescence in coastal waters
of the island of Laeso and along the Danish coast. Ctenophores cause
bioluminescence inside Limfjord and Mariagerfjord. Noctiluca attains
maximum abundance in the Belts and Kiel Bay in May and June. Lumines-
cence is very intense during August in the various bays around Kiel,
whereas in the open sea some distance from the coast it becomes in-
tense in September and October. Michaelis (1830) observed that the
blooming of Ceratium was most pronounced during windless, warm summer
days leading to some rather strong sheet-type displays in Kiel Bay.
Wintertime luminescence is rare, although sea light has been observed
under the ice at Kiel.

Very little bioluminescence occurs in the Baltic Sea. Tarasov
(1956) has indicated that some bioluminescence may be observed during
July and August in the western and central Baltic. Large numbers of
Pleurobrachia develop along the south coast of Finland in summer and
autumn and are probably responsible for some glowing-ball displays in
the Gulf of Finland. No significant bioluminescence has been reported
from the Gulf of Bothnia. The extremely low salinity would seem to pre-
clude the development of large numbers of luminous organisms.

SOUTHERN NORTH ATLANTIC AND SOUTH ATLANTIC OCEANS (FIGURES 2 and 3)

The percentages of total observations between 20°N and the Tropic
of Capricorn for three-month periods are as follows:

January through March ---------- 2T%
April through June ------------- 22%
July through September --------- 29%
October through December ------- 22%

15

The percentages of total observations for the three types of displays
are as follows:

Glowing-ball type ---------- 50%
Sheet type ----------------- 27%
Spark type ----------------- 23%

The various types of displays are quite evenly distributed through-
Out the year, although there are some exceptions. A maximum number of
glowing-ball and sheet-type displays occur from January through March,
whereas glowing-ball displays are least frequent from April through
June. Spark-type displays are least frequent from October through
December.

South of the Tropic of Capricorn, data are relatively meager.
Except for the shipping lanes from southwest Africa and southeast South
America, large regions of open ocean are seldom traversed by vessels.
Aithough intense displays probably occur south of this tropic, the
frequency of the different types of displays is not known; glowing-ball
displays probably predominate in the open ocean and sheet-type displays
near the coasts. Spark-type displays might be expected to occur more
often toward the south. The sparse data available appear to indicate
that displays occur evenly throughout the year in this vast southern
portion of the Atlantic.

In tropical waters containing the Equatorial currents, more bio-
luminescence has been observed east of the 30 W meridian, especially
toward the bulge of Africa, than west of this meridian. A number of
rather spectacular displays, principally glowing ball, have been ob-
served in this region. Near Sao Tome, observers on board the whale
catcher ENERM noted a display of blue light in which the wake glowed
and the sea surface appeared as if fireworks were going off. The dis-
play was due to Pyrosoma, millions of which covered the sea surface to
the horizon. The German research vessel METEOR, while sailing back and
forth across the Equator in December, January, and February 1926, noted
continuous bioluminescence in the water night after night. The prin-
cipal organism creating the light also was Pyrosoma, often in great
numbers.

Sheet-type displays occur most often in coastal regions, although
some strong displays have been observed in oceanic equatorial waters.
A typical sheet-type display was observed from the SS HERMINIUS in
December 1931 while sailing toward Capetown. The sea appeared com-
pletely “phosphorescent,” and it gave the "effect of the sea being
illuminated by some unseen light underneath." The vessel

"ran suddenly into a mass of pale, but intense green sea
with silvery-white edges to the waves, somewhat similar
in appearance to the illuminated dial of a wrist-watch,
when observed in the dark, only of much greater intensity.

16

9
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Fa oo

LEGEND
© JAN., FEB., MAR.
* APR., MAY, JUNE
EACH SYMBOL REPRE-
SENTS ONE DISPLAY
REPORT.

FIGURE 2 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, SOUTHERN
NORTH ATLANTIC AND SOUTH ATLANTIC OCEANS (JANUARY

THROUGH JUNE)
17

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The waves, breaking in the distance gave the appearance of
the sails of a small yacht having a beam of light played
upon them. As each bow wave broke, the white paintwork of
the bridge and vessel's superstructure was brilliantly
illuminated."

Sheet-type bioluminescence may resemble a "white sandy beach" in the
distance, as was noted off Rio Para, Brazil.

Numerous displays occur along the coast of Africa, being most pro-
nounced off the mouths of rivers. Discolored water from the Congo River
has been observed some 300 miles from shore, and bioluminescence from
rapidly developing bacteria and dinoflagellates may be seen through the
region affected by this river. The METEOR sailed through this region in
1926, and observers noted a diffuse luminescence all along the coast of
what was then French Equatorial Africa extending into the coastal waters
of Angola. The light was continuous and rather intense in the vicinity
of Mocamedes.

Farther south along the coast of southwest Africa, the incidence of
bioluminescence increases markedly. Here, coastal waters are very pro-
ductive because of abundant nutrients introduced by the upwelling
Benguela Current. Walvis Bay, which is representative of many bays
along this coast, is subject to periodic "red tides" and brilliant dis-
plays of bioluminescence. The "red tides," caused most frequently by
Noctiluca, are accompanied by mass mortalities of marine life. Asa
result, luminescent bacteria develop on the decaying organisms, and the
displays may range from dull to brilliant in the surface waters of the
bay.

In False Bay, Union of South Africa, exceptional displays have been
observed. One observer noted that the bay in November was covered by a
"creasy" froth, variously colored, that gave the water an unclean appear-
ance by day but caused it to resemble a lake of "molten gold" at night.
Strong bioluminescence occurs in Table Bay in December and January, when
dark red water develops.

In colder waters extending southward toward the Antarctic, an in-
ereased number of spark-type displays caused by euphausiids may be
expected throughout the year. The usual displays caused by various
jellyfishes and tunicates such as salps have been seen throughout the
extreme southern portion of the South Atlantic. Harbors of the Isla de
los Estados (Staten Island) often are filled in the early part of
December with medusae which are said to cause brilliant bioluminescence
at night. Salps cause some rather strong displays between southern
Africa and southern South America. Dinoflagellates occasionally are
abundant in waters near the Falkland Islands. Here the movements of
large fishes have been observed by their bioluminescent tracks.
Euphausiids have been observed luminescing both in the water and on
floating ice in February near the South Orkney Island.

We)

The incidence of bioluminescence increases northward along the east
coast of South America. Many luminescent forms are carried north by
the Falkland Current and develop in large numbers in regions such as
the convergence zone to the east of the Rio de la Plata.

Strong bioluminescence occurs within and seaward of the Rio de la
Plata. This region is one of the most bioluminescent parts of the South
Atlantic. Here, Darwin (1845) observed

"a sea that presented a wonderful and most beautiful spectacle
-»-.. the vessel drove before her bows two billows of liquid
phosphorus, and in her wake she was followed by a milky train.
As far as the eye reached the crest of every wave was bright,
and the sky above the horizon, from the reflected glare of

the livid flames, was not so utterly obscure as over the

vault of the heavens."

Commander Charles Wilkes (1845) on board the USS VINCENNES during the
famous exploring expedition of the U. S. Navy in 1838-42 observed in
this convergence zone "that the water was much discolored due to salps
during the day, and as night closed in the sea became very luminous,
the vessels passing through the water leaving long bright trains behind
them." Just after noting this luminosity, the ship encountered a tem-
perature drop in the water, probably due to the Falkland Current, but
the sea remained luminescent on each side of this front.

Caribbean Sea

More luminescence occurs in the eastern Caribbean than in the
western portion. Frequent displays occur along the coast of Venezuela,
principally in the Gulf of Paria and in the Golfo de Venezuela.

Exceptional bioluminescence may be observed in the bays located
along the coasts of various islands encompassing the Caribbean Sea.
Bahia Fosforescente (Phosphorescent Bay), located on the south coast
of Puerto Rico, is outstanding in this respect. In this bay sheet-type
bioluminescence occurs year round because of a continuously abundant
dinoflagellate community. The bay itself acts as a culturing vessel,
complete with the proper nutrients, the right temperature, and other
ecological factors needed to sustain a continuing rich community of
dinoflagellates. Bioluminescence is intense when the water is agitated
by boats or fishes. Bays, such as this, usually are bordered by man-
groves, with continual drainage from the land into the bay.

Oyster Bay, near Falmouth, Jamaica is another bioluminescent bay,
in which sheet-type bioluminescence occurs throughout the year. Harvey
(1952) visited the region and found the water "marvellously beautiful,
with every fish that moved outlined in fire and every wave looked as if
it were aflame." He indicated two responsible organisms, the marine
dinoflagellates Ceratium and Pyrodinium bahamense. Research concerning
bioluminescence and bioluminescent organisms was conducted in this bay

20

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LEGEND
+ JULY, AUG., SEPT.

beopett ty

* OCT., NOV., DEC.

EACH SYMBOL REPRE-
SENTS ONE DISPLAY
REPORT.

FIGURE 3 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, SOUTHERN

NORTH ATLANTIC AND SOUTH ATLANTIC OCEANS
(JULY THROUGH DECEMBER)

21

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by a joint Johns Hopkins University-U. S. Naval Oceanographic Office
team in February 1963. Further investigative efforts along this line
are to be carried out by this group from July 1965 to June 1966.

Until the early 20th century, another spectacularly bioluminescent
bay near Nassau, Bahama Islands was described regularly. However, the
environment of the bay was changed by the dredging of a channel, and the
water can no longer support large populations of luminescent organisms.

NORTHERN PACIFIC OCEAN AND ADJACENT SEAS (FIGURE 4)

Little information is available concerning the incidence of bio-
luminescence in the East China and Yellow Seas. Some displays occur in
the summer and autumn, when Noctiluca and Salpa often appear in very
large shoals; the former organism also may cause various degrees of
luminescence in coastal waters in spring. The water may appear greenish
and turbid by day because of masses of green-colored Noctiluca. This
particular species of Noctiluca has been observed to luminesce brightly
by Haneda (1955), although Ostrumoff (1924) and other workers consider
the green variety as nonluminescent. Peridinium and Ceratium (lumin-
escent dinoflagellates) have been observed in large concentrations near
Nagasaki in October.

Bioluminescence may be very brilliant at times along the west coast
of Kyushu and around some of its adjacent island groups. Observers on
the SS LAOMEDON noted that much reddish brown discolored scum was
floating in the water in April 1928 as they approached the Danjo Islands
from the southwest. After sundown the patches gave off a bluish green
glare. During the evening they observed long streaks of "phosphores-
cence"; the light from the individual streaks was so bright that lights
from passing ships were obscured by the glare. This display lasted
throughout the night. The streaks had an oily appearance indicative of
concentrations of Noctiluca, in this case most likely the pink colored
species.

The most prevalent type of bioluminescence in the Sea of Japan is
the sheet type, generally caused by pink Noctiluca. This dinoflagellate
is present throughout the year and is especially abundant in spring. It
and other luminescent dinoflagellates may cause a "red tide" along the
Japanese coast.

Luminescent dinoflagellates, particularly Peridinium, have been ob-
served in large reddish blooms during late summer along the Soviet coast
of the Sea of Japan. Luminescence in the water near Vladivostok is
caused by dinoflagellates from the latter part of August until the first
part of October. The light given off often is very intense. Euphausiids
or copepods contribute to the overall bioluminescence in August and
September. A diffuse luminescence consisting of sparkles often is noted
in bays such as the Bay of Patrokl (near Vladivostok) at this time, evi-
dence of the large number of euphausiids and copepods present in the
surface waters.

23

Spark-type bioluminescence caused by swarms of euphausiids also may
be observed in the Sea of Japan; it is blue and prevalent in. the northern
part. The light given off by these and other crustaceans is very intense
in April and is used by fishermen to detect moving herring schools.

Although reports of glowing-ball bioluminescence are lacking for the
Sea of Japan, this type of display probably occurs within the Tsushima
Current. Both Pyrosoma and Salpa often are observed in the Sea of Japan,
the former restricted to warm water regions in the south and east.
Haneda (1955) noted that numerous individuals of Beroe (a luminescent
etenophore) often were mixed with specimens of the luminescent squid
Watasenia scintillans in nets retrieved from the waters of Toyama Bay.

In the Sea of Okhotsk the spark type is the most commonly observed
bioluminescence. The displays are frequent from August through December
and often very intense, especially in September. According to Stukalin
(1934), unusually bright greenish white light may be observed in August
and September. The regions of cold water and the fronts of the cold
water masses appear to be the most favorable regions for the development
of many luminescent forms. Noctiluca occurs in large numbers in the sur-
face waters of the Gulf of Terpeniya and off southern Sakhalin in spring.
An influx of luminescent organisms, including ostracods, often occurs
along the north coast of Hokkaido and east coast of Sakhalin in Septem-
ber. Copepods such as Metridia are responsible for displays during May
and June.

Icebreakers in the Sea of Okhotsk have created some strong scintil-
lating green bioluminescence as they crashed their way through icefields
or bumped floating chunks. The light probably is given off by im-
prisoned luminescent organisms. These organisms occasionally make the
ice appear red, from which the term "bloody ice" has been derived.

Certain other luminescent displays, the shapes and movement of which
are unexplainable at this time, have been observed by observers in the
Sea of Okhotsk. These displays are best described as large expanding
circles of light which appear suddenly in the sea, or as waves of light,
or strips of light extending in various directions. Occasionally these
strips of light will rotate, resembling a "phosphorescent wheel" (see
section on this subject). Observers from the Sakhalin Branch of the
Russian Oceanological Institute on a research expedition noted such an
appearance (Priroda, 1956) 14 miles east of Cape Aniva in September 1953.
They reported as follows:

"On the night of the 9-10 of September unexpected bright
luminescence of the sea was observed across the stern of
the ship, which progressed rapidly, forming a rather large
ring in the center of which was the ship. It was very
brilliant near the ship, and was strongly luminescent as
it rapidly increased in size progressing from the ship to
the invisible horizon. The luminescence was intense and

2h

LEGEND
© JAN., FEB., MAR.
x APR., MAY, JUNE
+ JULY, AUG., SEPT.
* OCT.,NOV.,DEC.
EACH SYMBOL REPRE-
SENTS ONE DISPLAY
REPORT.

FIGURE 4 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, NORTH
PACIFIC OCEAN AND ADJACENT SEAS

25

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moving fast. The water did not contain organisms visible

to the eye. The intensity of the luminescence was sufficient
to read by. It seemed the rapidly moving waves of light were
accompanied by an unclear sound reminiscent of boiling water.”

Another sighting of a luminescent display was reported in an article
entitled "Huge Luminescent Mass Sighted in the Sea of Okhotsk" published
in the Sapporo Edition of the "Asahi Shimbun" in August 1964. The arti-
cle stated that just southeast of Cape'Terpeniya, Captain W. Takada of
the patrol ship TESHIO noted in August a spot of bright light which
measured a few meters in diameter at first and then expanded quickly to
400 meters in diameter; it then contracted to 30 meters in diameter and
disappeared into the sea in a whirling manner. The luminescence was as
strong as an electric lamp but not the same as would occur when a ship
broke seawater containing Noctiluca, for example.

Along the Pacific coast of Japan Noctiluca is particularly abundant
in spring, when it occurs in such great quantities at times that the
water, in addition to being discolored, becomes almost viscous in con-
sistency. This extraordinary abundance has been observed in Aomori Bay
and along the coasts of northeast Honshu and northeast Hokkaido. Spark-
type displays occur very frequently in late summer and early autumn in
the coastal regions as well as at some distance from the coast. These
displays predominate in the more northern waters of Japan, whereas
sheet-type and, to a lesser degree, glowing-ball-type bioluminescence
appear most frequently farther south. Very dark brown or blood-red
water caused by dinoflagellates has been observed in Tokyo Bay from
January through March, and the sheet-type bioluminescence produced is
brilliant at night. Many displays, often bright, have been observed in
the Gulf of Sagami where warm water organisms of the Kuroshio and cold
water organisms of the Oyashio are brought together.

Spectacular bioluminescence also has been observed during earthquakes.
"Fiery" columns of light were observed in the Gulf of Sagami during the
severe earthquake of 1923. The light was believed to have come from bio-
luminescent organisms violently stimulated by the strong shocks trans-
mitted through the water. Near Sanriku, Honshu the sea receded during a
tsunami. The exposed bottom was strongly luminescent with a bluish white
light of such strength that land objects were visible as if in daylight.
This display was attributed to the stimulation of the luminescent organ-
isms coating the bottom.

On the Pacific side of the Kuril Islands and to the north along the
east coast of Kamchatka, bioluminescent displays have been observed
throughout the year wherever the water is free of ice cover. The lumin-
escence is mostly sheet type and usually is seen in coastal waters in
the spring; spark-type bioluminescence also has been observed in the
spring, but generally some distance from the coast where large concen-
trations of luminescent euphausiids occur. Spark-type bioluminescence
is the most common type observed during the autumn and winter months.

27

Only meager data are available concerning bioluminescence in the
Bering Sea and northern part of the North Pacific Ocean. The few re-
ports available indicate that the majority of displays occur from April
through September, although the water probably is highly luminous at
times during autumn and winter. Noctiluca and other luminous dino-
flagellates are the principal light producers during summer, when
maximum bioluminescence apparently occurs; copepods and euphausiids
seem to cause the majority of displays during other seasons. Shoals of
luminescent salps and medusae have been reported in the warmer waters of
the North Pacific.

The maximum amount of bioluminescence in the region most probably
occurs in the zone where the Oyashio and Kuroshio meet. One observer,
while traveling across the front between the two currents, noted the sea
as a tremendous arc of "phosphorescence" which lighted up the ship as
though a green floodlight had been turned on.

In the western Aleutian Islands bioluminescence is pronounced during
summer and early autumn (July through October). Both sheet and spark
types have been observed during this period. The copepod Metridia has
caused strong greenish bioluminescence in the coastal waters of the
various islands of the Aleutians in the summer. Noctiluca reaches a
peak of abundance in late summer and is responsible for displays in the
surface waters off Attu, Kiska, and other islands.

Reports of bioluminescence are fragmentary in the northeastern part
of the North Pacific. However, bioluminescence undoubtedly is much less
frequent in the open ocean than near the coasts. Glowing-ball type
bioluminescence increases in frequency southward to the Hawaiian Islands.
Many species of luminescent medusae occur in the central Pacific.

Along the California coast many sheet-type displays occur during
spring and summer when red water is produced by large numbers of lumin-
escent dinoflagellates. Red water also may develop in winter when
unusually warm water conditions occur. Luminous water observed just
south of Monterey contained large numbers of dinoflagellates. The water
may be highly luminescent off San Diego and San Francisco in November.
This luminescence often is due to concentrations of salps or pyrosomes
and at times to ctenophores.

Along the coast of southern California and Baja California, fish
schools are located with the aid of bioluminescence. The kind of fish,
depth, and direction of movement all are revealed by the bioluminescence
ereated when they move through large concentrations of bioluminescent
organisms. Nets drawn through the water often retrieve a very luminous
slime made up of innumerable dinoflagellates.

28

EAST-CENTRAL AND SOUTH PACIFIC OCEAN (FIGURE 5)

Bioluminescence has been observed along the Pacific coast of Mexico
and Central America throughout the year. The majority of displays are
sheet type and glowing-ball type. Strong bioluminescence occurs in the
Gulf of California during autumn. Beebe (1942) observed bioluminescence
near Acapulco, Mexico during November. He described the display as "a
mass of boiling turquoise foam" extending behind the ship and noted "a
narrow intense curved line of pale green extending out from the side of
the ship as if a luminous serpent were gliding with us over the surface."
Brilliant displays of bioluminescence have been reported in Bahia Ballena,
Costa Rica in March.

In the western approaches to the Panama Canal, bioluminescence occurs
most frequently from January through June. The displays are primarily
sheet type and glowing-ball type. Unusual displays of crescent-shaped
"phosphorescence" and patches of pale green light often are reported.

The crescent-shaped light apparently was not affected by the movement of
a ship and most probably was caused by pulsations of the ship's engine,
similar to incidents reported in Asian waters. Bright balls of "phos-
phorescence" are commonly observed in February.

In the vicinity of the Galapagos Islands, marine life is rich and
includes a number of bioluminescent species. Beebe (1924) found the
marine life very distinct at night, each organism "glowing from a phos-
phorescent silhouette." Copepods and medusae are the main bioluminescent
forms in this region. Beebe also described concentrations of organisms
capable of causing luminescent displays that he observed between Cocos
Island and the Galapagos. He noted a gigantic current rip in April
caused by two westward flowing currents; at the juncture of the currents
the water was so full of plankton that it resembled soup. Single-celled
organisms capable of lighting up the sea at night also were reported by
him. Salps were present in great numbers. This enormous concentration
of plankton continued for several nights. Noctiluca glowed in the water
at night just south of Cocos Island in May.

The ocean off Peru and Chile within the Peru Coastal Current is
highly luminescent. However, marine life farther offshore in the Peru
(Humboldt) Current is meager, and only occasional displays of light have
been observed. Along the coasts of Peru and Chile, bioluminescence has
been known to occur most frequently from September through April.

The most intense sheet-type displays are said to occur along the
Peruvian coast from December through April, when "aguaje" (sick water)
develops in coastal waters. The water becomes greatly discolored
sometimes bright red or various shades of brown and often, because of
decomposing marine forms, gives off an offensive odor of hydrogen
sulfide. At night the bioluminescence is often extremely brilliant.
During periods when "El Nino" extends much farther south than normal,
as into northern Chilean waters, the water becomes highly discolored

29

LEGEND
© JAN., FEB., MAR.
* APR., MAY, JUNE
+ JULY, AUG., SEPT.
* OCT., NOV., DEC.
EACH SYMBOL REPRE-

SENTS ONE DISPLAY
REPORT.

FIGURE 5 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, EAST-
CENTRAL AND SOUTH PACIFIC OCEAN
30

because of massive blooms of luminescent and nonluminescent dino-
flagellates. Murphy (1926) refers to an observer who noted during an
"El Nino" disaster some years before that "after dark the sea broke out
in phosphorescent lightings all along the coast," and during the day
"the water was covered with blood-like patches many acres in extent."

One interesting report indicating a formation much like those ob-
served in the south Okhotsk Sea and southeast Asian waters was made by
Captain R. B. Bryant of the SS CURZCO sailing from Iquique, Chile to
Tocopilla, Chile in September 1960. He noted "very marked phosphor-
escence in the form of lines which appeared to be spinning around in
circular movement until dispersed by breaking waves." This appears to
be a form of "phosphorescent wheel."

In the South Pacific north of 60°S over twice as many displays
(about 69%) occur from September through January than during the rest
of the year. Most of the displays in this region are glowing-ball type,
usually produced by masses of salps, medusae, or Pyrosoma. A December
report noted "globules of phosphorescent matter that would brighten
momentarily when disturbed by the ship and cause sufficient light to
illuminate an area of the sea surface about 8 feet in diameter." It
also was observed that the breaking of waves some distance from the ship
was sufficient to cause the same reaction. Shoals of bright red
euphausiids often are noted in the southern regions of this area. The
shoals luminesce at night and cause a scintillating blue light, which
is quite striking to observers.

Bioluminescence probably occurs extensively throughout the year in
ocean waters near the various island groups of the South Pacific.
Although available data are inadequate, displays most likely occur with
equal frequency throughout the year.

One interesting phenomenon may be mentioned. In the Samoan Islands
the palolo worm swarms on the surface of the sea after dark in October
and November. This swarming takes place near coral reefs, and the water
is filled with vast numbers of threadlike worms which give out a pale-
green "phosphorescent light."

WESTERN SOUTH PACIFIC AND EASTERN INDIAN OCEANS (FIGURE 6)

The percentages of total observations to the north of the Tropic of
Capricorn for three-month periods are as follows:

January through March ------------ 16%
April through June --------------- 25%
July through September ----------- 39%
October through December --------- 20%

31

LEGEND
© JAN., FEB., MAR.
x APR., MAY, JUNE
+ JULY, AUG., SEPT.
© OCT.,NOV., DEC.
EACH SYMBOL REPRE-

SENTS ONE DISPLAY
REPORT.

FIGURE 6 SEASONAL DISTRIBUTION OF BIOLUMINESCENT DISPLAYS, WESTERN
SOUTH PACIFIC AND EASTERN INDIAN OCEANS
32

The percentages of total observations for the three types of displays
in this region are as follows:

Sheet. typel=- === === 57%
Spark type --------------- Th
Glowing-ball type -------- 36%

Fewer displays have been reported to the south of the Tropic of Capri-
corn. The percentages of total observations for this region for three-
month periods are as follows:

January through March ------------ 13%
April through June --------------- 34%
July through September ----------- 11%
October through December --------- hom

Data are inconclusive as to the types of displays most frequently ob-
served, although both sheet type and glowing-ball type bioluminescence
appear to be more frequent than the spark type.

Pyrosoma (the fire cylinder) is responsible for the more spectacular
glowing-ball displays throughout the year in this area. When luminescing,
as observers aboard the USS VANCE noted in September 1961 in the South-
west Pacific, they were visible at a distance of 75 to 100 yards.

Large shoals of medusae also create brilliant displays of light in
this area. In November 1950 observers on board the SS WAIHEMO noted a
brilliant "phosphorescent" region as they steamed along in the western
South Pacific. "Where the wash broke, the sea was brilliant blue, nearly
as bright and the same color as the are from a welding torch." Intense
patches of light about the size of dinner plates also were observed.

This reference to an arc light indicates the brilliance of some bio-
luminescence, sufficient to outline a ship in light.

Of special interest are the very brilliant snow white sheet-type
displays which occur in the Banda Sea and have been observed most fre-
quently during June through September. Because of these unusual
displays the Banda Sea often is referred to as the "white sea." A
characteristic of the displays is the suddenness with which they occur.
The sea, normal in appearance, suddenly becomes "milky" as if a light
switch had been thrown. The light is so strong that the sky may appear
to be lighted, and the ocean may give the appearance of a snow covered
land scene to observers. These exceptional sheet-type displays prob-
ably are due to large concentrations of dinoflagellates such as
Noctiluca.

An article in the Washington "Sunday Star" of 9 February 1958 con-

cerned recollections of an observer in the 1920's in Sandakan Harbor,
British North Borneo. He stated:

33

"Just after dark, as we looked towards shore, it seemed
that a river of fire was coming toward us. When we
entered it, it lit up the whole ship. As we passed
through it, the barnacles on the ship's bottom picked
up and retained some of the phosphorescent water. When
we were in dark water again, the bottom of the ship
glowed like an inverted candelabrum."

The luminous forms were undoubtedly dinoflagellates, such as Noctiluca.

In the South China Sea, Gulf of Thailand, Strait of Malacca, and
Andaman Sea bioluminescence is observed most frequently from April
through October. These displays are primarily the sheet and glowing-
ball types. Much discolored water occurs in these bodies of water, and
a high percentage of the plankton identified in the discolored portions
are luminescent forms. "Phosphorescent wheel" formations also occur in
these regions. Captain M. W. Lewsi, commanding the SS FENGTIEN, while
traveling in the South China Sea reported such an appearance in October
1961. He noted that a sample of water through which blue waves of
luminescence had passed was perfectly clear, indicating that the light
observed was given off by organisms not visible to the eye or that the
light produced was not biological in nature. Further discussion of this
particular aspect is contained in the section on "phosphorescent wheels.”

The coastal waters of the many islands of the Philippines are
strongly bioluminescent at times, usually because of concentrations of
luminescent dinoflagellates. Rust colored water is common in January
inside Manila Bay and is the result of masses of Peridinium, which
cause intense blue-green luminescence at night.

Immense numbers of Noctiluca develop in the vicinity of Hong Kong
from November through January, and the water is brilliantly luminescent
at this time. This condition also exists in other harbors and inlets
along the coast of south China and Hainan.

CENTRAL AND WESTERN INDIAN OCFAN AND ADJACENT SEAS (FIGURE 7)

Within the region north of the Equator the surface waters are highly
luminescent; the Arabian Sea is probably the most luminescent sea in the
world. The percentages of total observations in this region for three-
month periods are as follows:

January through March ---------- 23%
April through June! -=---===--=-— 12%
July through September --------- 48%
October through December ------- 17%

This distribution indicates the monsoonal influences on the movements of
surface waters, by which large numbers of bioluminescent organisms may

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drift together or by which upwelling increases surface nutrients and
thereby increases the production of marine forms capable of luminescing.

The percentages of total observations for the three types of dis-
plays in this region are as follows:

Sheet type -------------------- 75%
Spark type -------------------- 3%
Glowing-ball type ------------- 22%

The light given off by the sheet-type displays often is glaring in
intensity and much more spectacular than any reported from other ocean
areas of the world.

Some of the more unusual reports on surface bioluminescence are
from the Arabian Sea. Numerous patches of vivid bioluminescence were
noted on the surface of the sea by observers on the SS ELPHENOR in
September 1931. The patches seemed to be arising from the depths of
the ocean; the water swirled as if in an upward and spiral movement.
Later in the evening, it was found that a ray of light from a flash-
light could produce bioluminescence in the water wherever shown. Ob-
servers on board the USS DEVON in December 1929 noted small patches of
bioluminescence which suddenly expanded from diameters of about 6 feet
to diameters of 100 to 400 feet. In April 1956 observers on board the
MV DAGMAR SALEN noted a similar appearance to that of the crew of the
DEVON. The luminescence appeared to "bubble" up from below in the
form of a cone of light, which eventually spread out to a diameter of
100 to 150 years. Those on board the DAGMAR SALEN also observed the
water begin to "boil" because of thousands of mackerellike fish. One
authority believes this occurrence to be a rare occasion when the Deep
Seattering Layer broke through to the surface, as the mass appeared to
be made up of luminescent plankton and the fish feeding upon them.

A "milky sea" display was noted by observers on board the Greek
vessel SS IONNIS ZAFIRAKTS in August 1950 in the Arabian Sea. Second
Officer M. Mariakis reported that the horizon commenced to whiten as
the ship approached the luminous region, and then within the luminous
region the water assumed a milky color. The discoloration of the
water was considered different from "phosphorescence" by the reporter.
The phenomenon lasted about ys hours, and during that time the lower
layers of the atmosphere acquired a very thin whitish appearance,
reaching approximately 15° altitude and dimming the brilliance of the
stars. The sea was rough until the ship reached the luminous area,
where it became calm and glassy. This change may have been the result
of concentrations of oil-bearing luminescent organisms such as
Noctiluca. Captain A. Charasakis of the Greek vessel VERNICOS NICOLAS
in the Arabian Sea during August 1949 observed that the "whole surface
of the sea became brightly and deeply fluorescent" and "the surface
appeared as if crystalline in nature, through which a high tension
current of electricity passed, illuminating it brightly." Visibility

37

was hindered by the light given off. That this condition can continue
over a series of nights is indicated by the report of observers on
poard the SS PORT HUNTER during the nights of 18 - 21 August 1925.

"Extraordinary phosphorescence was noted in the sea.

The sea became luminous from horizon to horizon each evening
between 8:30 and 9:00 p.m., and disappeared in the morning
about an hour before daylight. It gave the effects of the
ship sailing in a sea of milk, at times very bright, and

it was possible to read the Azimuth Tables on the bridge

by the light, usually between the hours of 2 and Mb, Bseftihe

Bioluminescence in this area appears to occur much less frequently
south of the Equator than north of it. Reports south of the Equator
make up only about 10% of all those available north of 60°s. Aalal
three types of displays have been observed, the glowing-ball and spark
types of bioluminescence being predominant. Shoals of pyrosomes or
salps are responsible for most of the glowing-ball bioluminescence in
this region. The few reports available indicate that glowing-ball
bioluminescence has been observed most frequently in the period April
through June and again from October through December. Little is known
about the incidence of spark-type bioluminescence, but there is
probably an increased number of these displays in the direction of the
Antarctic Convergence, where euphausiids and copepods reach peak
abundance most likely in the same months as do the glowing-ball
organisms.

"PHOSPHORESCENT WHEEL" AND "WAVE" DISPLAYS (FIGURE 8)

Reports of 85 "phosphorescent wheel" displays and 26 "wave" dis-
plays (not plotted) are on file at the Oceanographic Office. The
category of "wave" displays includes such light formations as luminous
undulating waves, luminous rays extending through the water, and various
light patterns which cannot be categorized as "phosphorescent wheels."

Descriptive material concerning “phosphorescent wheel" and “wave"
formations is both voluminous and variable. The complexities encountered
in attempting to classify such appearances are illustrated by the follow-
ing two reports from "The Marine Observer."

MV SCOTTISH EAGLE. Captain R. R. Baxter. Bandar Maahur to
Kwinana. Observer, Mr. S. M. Grant, Chief Officer.

"23rd April, 1955, 1625 to 1635 G.M.T. When approaching
Jazirat Tunb Island a bright flashing light was observed
on the port bow, distant about 1 mile. Almost simultan-
eously another was observed on the starboard bow. On
approaching it was seen that these were two revolving
phosphorescent wheels. The ship passed between them,
the centres being about ; mile distant on either side.
The wheel on the port side appeared to revolve anti-
clockwise, and that on the starboard side clockwise.

38

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AV1dSIG ANO SILN&AS
“ddddd IOFWAS HOVA

39

The spokes of radius ¢ mile were from 6 to 12 feet broad at
the tips with about 15 feet between them. They passed with
a frequency of 1 $ sec. with a colour similar to that of a
dull electric light. Immediately we had passed these two
wheels, further phenomena were observed (eon, (On) the
port side concentric circles were seen to radiate from a
centre, with an effect similar to that of dropping some-
thing in still water. On the starboard side there appeared
lines, apparently moving away from the ship in a manner
similar to the wake. As the ship passed they gradually
faded from sight astern and had been in sight 10 or 15
minutes. At 1700 a similar phenomenon was observed to the
SE distant about 7 miles."

"The Master comments: 'The sky was overcast and the
atmosphere appeared to have more than the usual amount of
particles in suspension, but the all-round visibility was
very good. Jazirat Tunb Light was about 20 miles away
and its rotating beams were visible in the air throughout
their complete revolutions. Above the sea surface their
was apparently a layer of mist a yard deep.

"The bands of phosphorescent light appeared to float
on top of this layer, but on closer examination of the
beams as they passed vertically under the observer it
could be seen that the sea was affected to a considerable
depth. Each band was similar in colour and appearance to
the Milky Way, myriads of particles of light dust with
brighter and larger specks here and there. My impression,
especially during the concentric ring phenomenon, was of
shock waves causing the millions of organisms to light up
as the wave passed through them, then going dark until the
next wave struck them. I do not believe the organisms
themselves were on the move. The effect on the onlookers
seems to have been a feeling of weirdness, bordering on
fear, similar to that experienced by people ashore during
earthquake tremors.

"When the later distant phenomenon was observed it
made a glow on the horizon and could be made out with
binoculars but no shape or form. I believe this was
another group of wheels covering more than a square mile.'
Temperature, air 81°F, wet bulb 75.6°, sea 78°."

Position of the ship: 26°11'N, 54°55'R.

MV BRITISH EMPRESS. Captain A. Henney, O. B. E. Port
Okha to Persian Gulf. Observer, Mr. P. M. Alderton,
3rd Officer.

"Sth April 1953, 2125 Indian Standard Time. Commencing
from about NNW, shafts of pale white diffused light appeared,
apparently travelling on the surface of the water at a great
speed. Each shaft was several feet wide and they stretched
as far as the eye could see. At first they appeared in

ho

perfectly parallel lines, equally spaced, passing the ship at
about one every second, but after five minutes they wheeled
round in perfect formation and approached the ship from all
points of the compass. They came from only one compass point
at a time and each change of direction was swift and definite,
though not abrupt. The most frequent directions were from
NNW and SSE.

"After about 15 minutes the shafts occasionally formed
into a rotating radial movement in which they retained their
equal geometrical precision and the frequency of about one
per second. At this time the pattern was continually
changing about every 20-30 seconds from the parallel lines
to the wheel. The periods of transition were hardly notice-
able, but they were not abrupt. Each time the wheel appeared
it was in a different place. On one occasion there were two
distinct wheels visible at the same time. Throughout the
period the wheels appeared they varied in direction of
rotation, some clockwise and some anticlockwise. Five
minutes later the pattern became still more complicated
but remained perfectly regular and at 2150 the light faded
out over a period of 30 seconds.

"Although the light appeared to be on the surface of
the water it was completely unaffected by the wind and no
disturbance of the water was produced. The most notable
feature of the phenomenon was the effortless speed and
mathematical precision of movement. The only near analogy
I think of is that of being placed in the middle of a
large A scan when a large variable AC current is supplied.
The whole effect was one of great weirdness and errieness,
so much so that the look-out man came on to the bridge
quite scared, believing that he was suffering from hal-
lucinations.

"The ship's course was 290° (T), speed 10 kt and no

alteration of either took place during the observation.
The sea was that corresponding to Beaufort Scale wind
force 2-3, swell negligible, sea temperature 78 F. The
sky was cloudless, with perfect visibility, wind NW, force
2-3, air temperature 60 F."

Position of ship: 22°42'n, 68°03'E.

A simple "wheel" report appears to include the initial sighting of

a spot or pulsation of light on, above, or just under the sea surface,
sometimes observed at a great distance. This luminous spot usually
broadens and forms a pyrotechnic pinwheel effect, with long curving
luminous arms (spokes) that turn either clockwise or counterclockwise.
More than one wheel may appear, sometimes very small (less than a

meter in diameter) and at other times very large (covering the horizon).
The hub of such a wheel often is a glaring and intense white lumin-
escence, of such intensity at times that one observer noted it appeared
"like magnesium burning."

yl

The cause or causes of such phenomena are still unknown, although
the luminescence created appears to be related directly to luminous
organisms in the water. However, some theories have been advanced to
explain these formations. Kalle (1960) has suggested that these
appearances result from the action of submarine shock waves upon bio-
luminescent organisms. These shock waves, originating from submarine
earthquakes, impinge upon bioluminescent organisms and cause them to
light up. The light pattern perceived is due to the interference
pattern created by shock waves and reflected waves. When in phase,
the luminescence formed is intense; when out of phase, it is weak.
Most of these formations have been observed in regions of shallow
water where such interference patterns can occur. According to Kalle,
the pattern created appears as a wheel or some closely allied appear-
ance. He also indicates that "exploding" luminescence, noted most
often in deep waters, is due to the arrival of submarine shock waves
at the surface. The pattern occurring resembles a bubble of light
which rises from the depths, bursts at the surface, and rapidly ex-
pands into a large circle of luminescence.

Nicol (1961) expressed doubts about this theory, giving as his
reasons the patchiness of the distribution of bioluminescent organisms
and the fatigue experienced by these organisms after continual stim-
ulation. Both the symmetry and duration of the pattern would be
greatly affected, the wheel and wave would be broken in appearance,
and the intensity would gradually lessen after each shock. This
occurrence has not been reported.

Hilder (1962) advanced the theory that "phosphorescent wheels"
were caused by electromagnetic waves, such as would occur in magnetic
fields. The distortion of the earth's magnetic field caused by a ship
moving through a region, magnetic anomalies created by sunspots, and
the ultrahigh frequency of radar are all implicated by him in the
development of wheels or waves. He reported an effect in the Solomon
Islands area in March 1955 in which luminescence of the sea was a
function of the radar emission. When the radar was turned on, the
luminescence appeared; when the radar was turned off, the luminescence
weakened considerably. Except for one other report, no other obser-
vations, even during conditions of total blackout, have been reported
concerning the above phenomenon. WNicol's argument is relevant to this
theory also.

Both light and sound waves have been implicated in the stimulation
of bioluminescence. The sound emitted by ship engines has been corre-
lated with the pulsations of light observed, also, observers have noted
that a beam of light on the surface of the sea activated biolumines-
cence. A bright green streak may be noted as the beam from a flash-
light is moved across the surface of the sea. When the light is
turned off, the bioluminescence disappears after a few seconds. In
still other situations, pinpoints of light, much like the end of a
lighted cigarette, flashed on when a light beam passed over and
immediately disappeared as the light was removed.

2

Although most investigators believe that the luminescence during
"wheel" displays is produced by marine organisms in the water, it is
possible that some other type of luminescence may be occurring, if not
exclusively, at least as an adjunct to the bioluminescence. Of the
displays reported (Figure 7), most occurred in the following areas:
southern Persian Gulf, Strait of Hormuz, seaward of the Mouths of the
Indus, Strait of Malacca, Gulf of Thailand, and southern South China
Sea. Displays occurred to a lesser extent in the northern Andaman Sea
and along the north coast of Java. All of these regions are shallow,
and rather extensive blooms of phytoplankton occur in all quite
regularly. The water often becomes supersaturated with oxygen during
these blooms.

Strong pressure waves passing through a solution of high gas con-
tent may cause an electroluminescenee due to the breakdown of bubbles
of gas within the solution. It is possible that this phenomenon may
occur in the sea also. Shock waves (from submarine earthquakes or
underwater volcanic activity), which often are strong enough to jolt
a ship, might also be strong enough to cause luminescence in sea water
containing a very high gas content such as may occur in this area.

Although the occurrence of "phosphorescent wheels" or "waves" is
unpredictable at present and the reports are not numerous, the avail-
able records indicate that these phenomena are more frequent during
particular months in four areas. In the southern Persian Gulf-Strait
of Hormuz area about 67 percent of reported displays have occurred
from April through June. Off the Mouths of the Indus and in the Strait
of Malacca approximately 75 percent of reported displays have occurred
from July through December, and in the Gulf of Siam and the southern
South China Sea 75 percent of reported displays have occurred from
April through June and from September through December.

POLAR REGIONS

Although little information is available concerning biolumines-
cence in the ocean surrounding Antarctica, some rather strong spark-
type displays have occurred along the fringe of the Antarctic Con-
vergence. Here massive concentrations of euphausiids have been noted.
The light they give off may be intense at times but more often is weak.

More bioluminescence has been observed in the ocean areas adjacent
to the Arctic Ocean, but information still is meager. Dinoflagellates
produce luminescence in the waters around Svalbard (Spitsbergen) ;
crustaceans also may create varying degrees of bioluminescence near
these islands. Tarasov (1956) reported an observation by K. S.
Badiginin in January 1940 at 81°911'N, 3°50'E of bright greenish
" phosphorescence" lighting up the waves of an ice-filled ocean, rem-
iniscent of light observed in the Indian Ocean at a previous time.

43

Snow and ice sometimes luminesce in these colder regions as a
result of entrapped dinoflagellates and crustaceans. One observer noted
his luminescent footprints in the snow as he walked along the edge of
the sea after dark. He described very intense but quickly fading bluish
white light outlining his footprints. Later investigations attributed
this light to luminescent crustaceans washed in from the sea into the
snow.

Tsachenko (1914) stated that luminescence was a common phenomenon
near Murmansk during autumn nights. Although the type of biolumines-
cence was not mentioned, certain luminescent dinoflagellates appear to
reach a maximum development in the waters of the western Barents Sea in
November and December. Other luminescent dinoflagellates are most abun-
dant in August. All of the seas fringing the Arctic Ocean probably
experience some spark-type bioluminescence due to crustaceans.

BIBLIOGRAPHY

Agassiz, A. 1888. Three Cruises of the United States Coast and
Geodetic Survey Steamer BLAKE. Houghton, Mifflin and Co.
Boston and New York. The Riverside Press, Cambridge, Mass.

Viel, i. Sh i.

ASAHI Newspapers, Sapporo Edition. August 13, 1964. Translated
article entitled "Huge Luminescent Mass Sighted in the Sea of
Okhotsk."

Beebe, W. 1924. Galapagos: World's End. Putnam and Sons. New
York and London. L29 C

---- 1942. Book of Bays. Harcourt, Brace and Co. New York. 302 pp.

Boden, B. P. and E. M. Kampa. 1964. Planktonic Bioluminescence.
Oceanography and Marine Biology Annual Review. H. Barnes, Editor.

George Allen and Unwin Ltd. London. Vol. 2. pp. 341-371.

Clark, W. D. 1963. Function of Bioluminescence in Mesopelagic
Organisms. Nature Vol. 198. No. 4887. June 29, 1963. pp.
12hh-12h6.

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46

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h7

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Hydrographic Office.

48

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Security Classification

DOCUMENT CONTROL DATA - R&D

(Security classification of title, body of abstract and indexing annotation must be entered when the overall report is classified)
Os ei Ce aS a
Unclassified
U.S. Naval Oceanographic Office ACESS

3. REPORT TITLE

THE DISTRIBUTION AND CHARACTERISTICS OF SURFACE BIOLUMINESCENCE IN THE OCEANS

4. DESCRIPTIVE NOTES (Type of report and inclusive dates)

Final report

5. AUTHOR(S) (Last name, first name, initial)

Staples, Robert F.

6- REPORT DATE Ja. TOTAL NO. OF PAGES 7b. NO. OF REFS
March 1966 oh 5

Ba. CONTRACT OR GRANT NO. 9a. ORIGINATOR'S REPORT NUMBER(S)

TR-18)

6. PROJECT NO.

9b. OTHER REPORT NO(S) (Any other numbers that may be assigned
this report)

10. AVAILABILITY/LIMITATION NOTICES

All distribution of this publication is controlled. Qualified DDC users shall
submit requests for copies to the: Commander, U.S. Naval Oceanographic Office,
W; hington D TTN: Code

ile Sra EMENTARY NOTES : 12. SPONSORING MILITARY ACTIVITY
U.S. Naval Oceanographic Office
Washington, D.C. 20390

13. ABSTRACT

Approximately 3,000 reports of bioluminescent displays on file
at the U.S. Naval Oceanographic Office are charted by three-month
periods. In addition, various aspects of bioluminescence, biolumine-
scent organisms, seasonal and geographic distribution, and displays
are discussed, A serarate chart and a discussion of the puzzling
phenomena of the "phosphorescent wheel" and "wave" displays are
included, This report is published as part of the Oceanographic
Office program in marine biology.

DD. 5.1473 Unclassified

Security Classification

Unclassified

Security Classification

KEY WORDS

Bioluminescence
Bioluminscent displays
Bioluminescent organisms

"bhosphorescent wheel" and "wave" displays

"burning sea"
"Milky sea"
luminescent sea
winter ocean
the white water

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