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DEPARTMENT OF TERRESTRIAL MAGNETISM

J. A. Fleming, Director

Scientific Results of Cruise VII of the Carnegie during 1928-1929
under Command of Captain J. P. Ault

BIOLOGY- V

The Genus Ceratium in the Pacific
and North Atlantic Oceans

HERBERT W. GRAHAM
NATALIA BRONIKOVSKY

CARNEGIE INSTITUTION OF WASHINGTON PUBLICATION 565

WASHINGTON, D. C.
1944

This book first issued December 30, 1944

PREFACE

Of the 110,000 nautical miles planned for the seventh
cruise of the nonmagnetic ship Carnegie of the Carnegie
Institution of Washington, nearly one -half had been com-
pleted on her arrival at Apia, November 28, 1929. The
extensive program of observation in terrestrial magnet-
ism, terrestrial electricity, chemical oceanography,
physical oceanography, marine biology, and marine me-
teorology was being carried out in virtually every detail.
Practical techniques and instrumental appliances for
oceanographic work on a sailing vessel had been most
successfully developed by Captain J. P. Ault, master and
chief of the scientific personnel, and his colleagues. The
high standards established under the energetic and re-
sourceful leadership of Dr. Louis A. Bauer and his co-
workers were maintained, and the achievements which
had marked the previous work of the Carnegie extended.

But this cruise was tragically the last of the seven
great adventures represented by the world cruises of the
vessel. Early in the afternoon of November 29, 1929,
while she was in the harbor at Apia completing the storage
of 2000 gallons of gasoline, there was an explosion as a
result of which Captain Ault and cabin boy Anthony Kolar
lost their lives, five officers and seamen were injured,
and the vessel with all her equipment was destroyed.

In 376 days at sea nearly 45,000 nautical miles had
been covered (see map, p. iv). In addition to the exten-
sive magnetic and atmospheric-electric observations, a
great number of data and marine collections had been
obtained in the field of chemistry, physics, and biology,
including bottom samples and depth determinations.
These observations were made at 162 stations, at an av-
erage distance apart of 300 nautical miles. The distri-
bution of these stations is shown in the map, which de-
lineates also the course followed by the vessel from
Washington, May 1, 1928, to Apia, November 28, 1929.
At each station, salinities and temperatures were ob-
tained at depths of 0, 5, 25, 50, 75, 100, 200, 300, 400,
500, 700, 1000, 1500, etc., meters, down to the bottom or
to a maximum of 6000 meters, and complete physical and
chemical determinations were made. Biological sam-
ples to the number of 1014 were obtained both by net and
by pump, usually at 0, 50, and 100 meters. Numerous
physical and chemical data were obtained at the surface.
Sonic depths were determined at 1500 points and bottom
samples were obtained at 87 points. Since, in accord-
ance with the established policy of the Department of
Terrestrial Magnetism, all observational data and ma-
terials were forwarded regularly to Washington from
each port of call, the records of only one observation
were lost with the ship, namely, a depth determination
on the short leg between Pago Pago and Apia.

The compilations of, and reports on, the scientific
results obtained during this last cruise of the Carnegie
are being published under the classifications Physical
Oceanography, Chemical Oceanography, Meteorology,
and Biology, in a series numbered, xmder each subject,
1, II, and in, etc.

A general account of the expedition has been prepared
and published by J. Harland Paul, ship's surgeon and ob-
server, under the title The last cruise of the Carnegie ,
and contains a brief chapter on the previous cruises of
the Carnegie , a description of the vessel and her equip-
ment, and a full narrative of the cruise (Baltimore, Wil-
liams and Wilkins Company, 1932; xiii + 331 pages with
198 illustraUons).

The preparations for, and the realization of, the pro-
gram would have been impossible without the generous
cooperation, expert advice, and contributions of special
equipment and books received on all sides from inter-
ested organizations and investigators both in America
and in Europe. Among these, the Carnegie Institution of
Washington is indebted to the following: the United States
Navy Department, including particularly its Hydrographic
Office and Naval Research Laboratory; the Signal Corps
and the Air Corps of the War Department; the National
Museum, the Bureau of Fisheries, the Weather Bureau,
the Coast Guard, and the Coast and Geodetic Survey; the
Scripps Institution of Oceanography of the University of
California; the Museum of Comparative Zoology of Har-
vard University; the School of Geography of Clark Uni-
versity; the American Radio Relay League; the Geophys-
ical Institute, Bergen, Norway; the Marine Biological
Association of the United Kingdom, Plymouth, England;
the German Atlantic Expedition of the Meteor , Institut
fUr Meereskimde, Berlin, Germany; the British Admiral-
ty, London, England; the Carlsberg Laboratorium, Bu-
reau International pour I'Exploration de la Mer, and
LaboratoireHydrographique, Copenhagen, Denmark; and
many others. Dr. H. U. Sverdrup, now Director of the
Scripps Institution of Oceanography of the University of
California, at La Jolla, California, who was then a Re-
search Associate of the Carnegie Institution of Washing-
ton at the Geophysical Institute at Bergen, Norway, was
consulting oceanographer and physicist.

In summarizing an enterprise such as the magnetic,
electric, and oceanographic surveys of the Carnegie and
of her predecessor the Galilee , which covered a quar-
ter of a century, and which required cooperative effort
and unselfish interest on the part of many skilled scien-
tists, it is impossible to allocate full and appropriate
credit. Captain W. J. Peters laid the broad foundation of
the work during the early cruises of both vessels, and
Captain J. P. Ault, who had had the good fortune to serve
under him, continued and developed that which Captain
Peters had so well begun. The original plan of the work
was envisioned by L. A. Bauer, the first Director of the
Department of Terrestrial Magnetism, Carnegie Institu-
tion of Washington; the development of suitable methods
and apparatus was the result of the painstaking efforts of
his co-workers at Washington. Truly, as was stated by
Captain Ault in an address during the commemorative
exercises held on board the Carnegie in San Francisco,
August 26, 1929, "The story of individual endeavor and
enterprise, of invention and accomplishment, cannot be
told."

Dr. H. W. Graham, who succeeded H. R. Seiwell as
chemist and biologist, had charge of the biological work
on board the Carnegie from August 1929 until the loss of
the vessel at Apia, Samoa. After his return to this coun-
try. Dr. Graham was placed in charge of the biological
collections, attending to their subsequent care, segrega-
tion, and distribution to various specialists for examina-
tion and report, he himself undertaking the reporting of
the Peridineae (Dinoflagellata). His memoir, "Studies
in the morphology, taxonomy, and ecology of the Peri-
diniales," is Biology-Hi of this series and in it Cerati-
um is briefly discussed. He also examined and prepared
a report on the "Phytoplankton," which is the first of
twelve biological reports in Biology-IV.

The present report discusses a single peridinian--

iii

PREFACE

genus Ceratium of the family Ceratiaceae. Dr. Graham
was assisted in the laboratory work necessary for the
present volume by Mrs. Natalia Bronikovsky, who did
the careful microscopic work which the dinoflagellate
investigations demand.

The genus Ceratium is distributed over all the oceans
of the world and is one of the most valuable genera of the
peridinians for distributional studies. Not only are there
cold- and warm-water species, but many species show
minor phenotypic variations which are useful in tracing
dynamic conditions. Fifty-eight species of this genus
were found in the Carnegie material and are discussed
here. Distributional and environmental data for these
different species are given in the appendix tables (pp.
47-161) and charts (pp. 187-207) whenever such infor-
mation is available.

The area traversed by the Carnegie was divided into
five regions on the basis of Ceratium floras (chart 45).
Two of these regions are in the North Atlantic and three
are in the Pacific Ocean. They are also characterized
by particular hydrographic conditions.

This manuscript was completed by Dr. Graham in
1938. Thus some papers printed since then are not con-
sidered.

The present volume is the eleventh in the series
"Scientific results of cruise Vn of the Carnegie during
1928-1929 under the command of Captain J. P. Ault."
This is the fifth of the Biological Reports.

J. A. Fleming
Director, Department of Terrestrial Magnetism

CONTENTS

Page

Introduction 1

Hydrography of Regions Investigated 1

Currents 1

Temperature 2

Salinity 2

Phosphate 3

Horizontal Distribution of Ceratlum . 3

Ceratium Life Zones in the Areas Traversed by

the Carnegie 3

Richness of Species in Different Areas 6

Factors Affecting the Horizontal Distribution of

Ceratium 7

Salinity 7

Nutrient Salts 7

Temperature 7

Currents 8

Other Factors 10

Page

Comparison of the Atlantic and Pacific Waters on

the Basis of Their Ceratium Floras 10

The Vertical Distribution of Ceratium 12

The Ceratium Species of the Carnegie Collection . 14

Subgenus Poroceratium 14

Subgenus Blceratium 16

Subgenus Amphiceratium 22

Subgenus Euceratium 25

Literature Cited 46

Appendix Tables 1-54 47

Figures 1-27 163

Charts 1-54 187

Index 209

/v

IVl;

vii

THE GENUS CERATIUM IN THE PACIFIC AND
NORTH ATLANTIC OCEANS

INTRODUCTION

The present paper, which constitutes a part of the
Carnegie plankton reports, deals only with the one peri-
dinian — (dinoflagellate) genus Ceratium. This genus
consists of a large number of species of which many are
very common and widespread marine forms. They con-
stitute a characteristic and often dominant part of the
planktonic community. The present report deals with
fifty-eight species, which comprise all but one or two of
the known marine forms as well as three new species.

Because of their constant participation in the ma-
rine phytoplankton, the Ceratia have of late been given
considerable attention in general oceanographic re-
searches. Thus, Peters (1934) has investigated the dis-
tribution and biology of the Ceratia collected by the Me-
teor during its Intensive oceanographic investigation of
the South Atlantic Ocean in 1925 to 1927 and Nielsen
(1934) has done likewise for the collections of the Dana
made in the SouthPacific during 1928 to 1930. The pres-
ent collections are unique in their extensive nature,
bringing together comparable material from widely sep-
arated regions of the Pacific and North Atlantic oceans.

In the Atlantic the stations extended along the Gulf

Stream or North Atlantic West Wind Drift, in the North
Sea, in the arctic currents about Iceland and Greenland,
and In the Labrador Current. There is also a line ex-
tending through the Sargasso Sea to the North Equatorial
Current and Equatorial Countercurrent. From here
there are stations running along the former current and
into the Caribbean Sea.

In the Pacific the collections are even more exten-
sive. There are two lines crossing the equatorial cur-
rents; one in the central Pacific between longitudes 140°
and 150° west, the other west of this at about longitude
175° west. Tropical stations in the North Pacific are
scattered widely over the east and west parts; north of
the Marshall Islands, between Guam and Japan; between
the United States and Hawaii; and north and south of Ha-
waii. In the South Pacific there is a line of stations in a
region of indefinite currents between Peru and Samoa.
There are stations from Peru southeastward at 40° south
latitude and north to Easter Island. Other stations are
in the region of the Galapagos Islands and the Panamic
area.

THE HYDROGRAPHY OF THE REGIONS INVESTIGATED

Currents

North Atlantic Ocean. --The central part of the North
Atlantic s»irface water consists of a huge vortex running
clockwise. The southern part of this circuit is repre-
sented by the North Equatorial Current which has a west-
erly course and is composed of water of high temperature
and low nutrient content. This current merges with the
Florida Current which has a northerly set off the east
coast of the United States. It, too, is composed of warm,
nutrient-poor water. This current, in turn, becomes the
North Atlantic West Wind Drift which flows eastward to
the coast of Europe. This current loses its tropical
characteristics as it progresses, becoming colder and
richer in nutrient salts. When it reaches Europe, some
of this water is deflected to the south to join the North
Equatorial Current, and thus to resume the circuit;
other parts continue northeastward around the British
Isles and finally into the Arctic Ocean. To the north of
the West Wind Drift there -is a general movement of arc-
tic water southward, more specifically in the form of the
Greenland Current and the Labrador Cvirrent. This cold,
nutrient-rich water meets the warm water to the south
at the polar front (subpolar convergence) which lies at
about latitude 40° north off the coast of America, and
from there extends west northwestward (see chart 50).
The surface temperatures at this convergence drop sud-
denly from 21° to 11° C. K should also be noted that

within the North Atlantic West Wind Drift there is a
sudden drop in temperature and increase in nutrients at
about 40° west longitude. Although the Carnegie stations
are rather widely separated in this region, the change is
striking, with a surface temperature of 21.°18 recorded
at station 14 and 15.°50 at station 3. Similarly the phos-
phate content of the upper 50 meters increased from 14
to 64 mg P04/m3. In addition to the above stations there
were four stations in the Caribbean Sea. The water here
was the warmest of any Atlantic region visited, with tem-
peratures all above 28° C, and it was practically devoid
of phosphate.

North Pacific Ocean . - -The main body of the North
Pacific is involved in a clockwise rotation similar to that
in the North Atlantic although it is modified by the differ-
ent configuration of the continents. The North Equatorial
Current is well developed; itsmiddle part is about at 10°
north latitude. It is warm, poor in nutrients, and has
temperatures as high as 29.°5 C. This current turns
northward off the Asiatic coast where it is known as the
Kuroshio Current, just as the corresponding current of
the Atlantic becomes the Florida Current. The Kuroshio
is warm and poor In nutrients. The waters of this cur-
rent turn eastward and flow south of the Aleutian Islands
as the North Pacific West Wind Drift. The waters now,
howevei , are cold and rich in nutrients, with surface
temperatures around 8° C, and the phosphate content of
the upper 50 meters is over 100 mg P04/ni3. These

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

waters are deflected to the south by the continent of
North America, along the west coast of which they flow
as the California Current. By the time this water has
reached a latitude of 30° north it has attained tropical
temperatures (above 20° C) and has become depleted of
nutrient salts. It continues into the North Equatorial
Current to complete the circuit. The polar front or
subpolar convergence of the North Pacific extends north-
eastward from Japan at a point at about 38° north lati-
tude. At this line the cold arctic waters of the Oyashio
meet the warm waters of the Kuroshio. Not only does
the North Pacific Intermediate Current originate here,
but there is considerable mixing of the two types of wa-
ter at the surface, so that from here the water of the
Kuroshio or the North Pacific West Wind Drift is colder
and richer in nutrients because of the admixture of cold
and rich water from the north. All the Carnegie stations
off Japan were to the south or east of the line of conver-
gence as indicated by Schott (1935), but in the hydro-
graphic studies of the Carnegie (see Sverdrup, 1944)
there was found in the upper 500 meters a definite line
of demarcation between the cold water to the north and
the warm water to the south, principally at the latitude
of 38° north, between stations 115 and 116.

A loop of Carnegie stations, with a center about
northeast of Hawaii, lies inside the main current system
of the North Pacific, and extends into the subtropical
convergence. This central part of the North Pacific can
be likened to the Sargasso Sea of the Pacific. The water
is of high temperature and low nutrient content to com-
paratively great depths. For instance, the temperature
is often above 20° C to more than 100-meters depth with
total depletion of phosphate.

South Pacific Ocean. --In the South Pacific also, the
current system is in the form of an extensive vortex; in
this case, however, it rotates in a counterclockwise di-
rection since it is in the Southern Hemisphere. In the
south of this circuit there is the South West Wind Drift
which travels easterly around the earth in a more or
less uninterrupted manner in the subantarctic latitudes.
This current is cold and rich in nutrients. It lies a little
to the south of the most southerly Carnegie station, sta-
tion 60, at 40° south. An offshoot of this great oceanic
stream, the Humboldt Current, extends northward up the
west coast of South America and brings cold, rich water
into this area. The waters from this region turn to the
left and flow westward as the South Equatorial Current.
The main part of this current lies at about 3° north lati-
tude. The water of this current increases in tempera-
ture and decreases in nutrient content as it progresses,
so that in the western Pacific it is very warm and poor
in nutrients, whereas in the eastern part, near its ori-
gin, it is not so warm (temperatures of 20° or less
around the Galapagos) and is richer in nutrients (25 to
50 mg P04/m3). It should be noted, however, that the
waters of the South Pacific are nowhere depleted of
phosphate although the surface temperatures may be
several degrees higher than in depleted areas of the
North Pacific. The South Equatorial Current gives off
numerous, If not continuous, offshoots to the south. One
of these offshoots departs at about longitude 120° west,
and thus forms a minor vortex with the Humboldt Cucr-
rent which has been called the Easter Island Eddy by
some. The water in this eddy showed the lowest phos-
phate content of any water investigated by the Carnegie
in the South Pacific. The currents of the southwestern
Pacific will not be discussed as no Carnegie stations

were occupied in that area. The Equatorial Countercur-
rent is a narrow, shallow stream extending eastward in
a direction opposite to its neighbors, at about latitude 6°
north. It is warm and poor in nutrients to the west, but
has lower temperatures and higher nutrient content to
the east, probably because of admixture with water from
the southern current system.

There are also Carnegie stations in the Panamic
area. The water in this region probably has a multiple
origin, a has both a high temperature and a high nutri-
ent content.

Temperature

In the North Atlantic south of the subpolar conver-
gence, the surface temperatures are mostly above 20° C
(chart 50). There is an increase in temperature south-
westward to maxima above 28° in the Caribbean region.
In the North Atlantic West Wind Drift the temperatures
rapidly decrease to 11° and 12° at Europe. At the other
stations, farther north, the surface temperatures are
mostly around 10° C.

In the North Pacific there are six stations in water
with surface temperatures less than 10° (stations 119 to
124) south of the Aleutian Islands to the Kurils. Toward
Japan there is a rapid increase in temperature south-
ward so that there are only three stations in the transi-
tion zone between the cold northern water and the warm
tropical water of the south, fii the eastern Pacific, on
the other hand, this transition is very gradual so there
are seven widely spaced stations between the 10° water
south of the Aleutians and the 20° water of the tropics,
stations 125 to 131.

to the tropical regions of the Pacific the surface
temperatures are mostly between 22° and 28° with the
highest temperatures in the west. Temperatures of less
than 20°, however, were recorded at two stations in the
region of the Galapagos, stations 42 and 43. The effects
of the Himiboldt Stream are noted in a general lowering
of the temperatures in the southeastern Pacific region,
although temperatures as high as 25.°27 were recorded
off Peru. The Carnegie stations south of 20° south lati-
tude were so far from South America that they do not
show the severe effects of this current. Only three of
the stations showed temperatures less than 20°. Farther
south, however, beyond 30° south, the effects of the South
West Wind Drift are very noticeable, with a fairly rapid
reduction of the surface temperatures to 14.°97 at 40°
south at station 60.

Salinity

The central North Atlantic in the Sargasso Sea re-
gion has the highest surface salinities of any area trav-
ersed by the Carnegie, namely over 37 per mille (chart
51). In the North Equatorial Current the salinity is
about 36 per mille; at the cold northern stations it is
mostly around 35 per mille, although there was one sta-
tion southeast of Newfoundland where the salinity was as
low as 32.66 per mille.

In the central North Pacific the surface salinities
are slightly more than 35 per mille. At the subpolar
convergence this is reduced to less than 33 per mille
and the salinities of the North Pacific West Wind Drift
remain under this value to northern California. In the
Panamic region the salinity is quite low, namely, 30 per
mille. The values increase westward from this area in

HORIZONTAL DISTRIBUTION OF CERATIUM

the North Equatorial Current with a tongue of 34 per
mllle water reaching 140° west longitude.

fii the South Pacific there is a zone of maximum sa-
linity centered at 20° south latitude and about 125° west
longitude with values over 36 per mille. Salinities de-
crease westward to 35 per mille at Samoa. Southeast-
ward from the zone of maximum the surface salinities
decrease to 34 per mille.

Phosphate

Since the nutrient salt content of sea water is known
to be important in the total production of the phytoplank-
ton, it Is necessary to examine the variation in nutrients
as a possible important factor In the distribution of in-
dividual species. Phosphorus and nitrogen are the most
important plant foods in this respect. Only phosphate
data are available for a discussion of the oceans as a
whole but, since the variations in quantities of phosphate
and nitrate run parallel, a description of phosphate dis-
tribution will suffice to give a general picture of the dis-
tribution of phytoplankton foods in the areas investigated
by the Carnegie . The discussion that follows deals with
values which are the means of observed values for the
surface and for 50 meters, from Carnegie data.

The warm North Atlantic waters are practically de-
void of phosphate to 40° north latitude. Two stations just
south of this latitude (stations 1 and 2) showed values
above 10 mg P04/m3. North of 40° north latitude the
mean values for the upper 50 meters were practically
everywhere between 25 and 75 mg P04/m3.

Most of the great area of the North Pacific between
latitudes 10° and 38° north is practically devoid of phos-
phate, with values less than 10 mg. Along the subpolar
convergence northeast of Japan the phosphate content
rapidly increases northward to over 100 mg. In the
North Pacific West Wind Drift the values are well above
100 mg to longitude 150° west where they gradually be-
gin to drop off southward. This southward gradient is in
the California Current. Close off California the values
are between 25 and 75 mg. Farther out there is a rapid
decrease to the very low values of the "Sargasso Sea of
the Pacific." The North Equatorial Current lies in this
area of extremely low phosphate content.

Nowhere to the south of this current in the Pacific
is there an area devoid of phosphate. The area of lowest
phosphate content in the South Pacific is in the Easter
Island Eddy. At only two stations here, however, did the
values fall below 10 mg. The subantarctic waters of the
South Pacific West Wind Drift are rich in nutrient salts.
The effects of this are shown by phosphate values above
50 mg at Carnegie stations 60 and 61 at about 40° south
latitude. Values over 100 mg were found along the Pe-
ruvian coast, probably the combined effect of the Hum-
boldt Current and local upwelling. In the general region
between Peru and the Galapagos and in the Panamic re-
gion the values are above 50 mg. In the South Equatorial
Current and its offshoots to the south the mean phosphate
values for the upper 50 meters are mostly between 25
and 50 mg as far west as Samoa. South and west of here
there is a tendency for the values to fall off.

HORIZONTAL DISTRIBUTION OF CERATIUM

Ceratium Life Zones in the Areas
Traversed by the Carnegie

The area traversed by the Carnegie can be divided
into five regions on the basis of the Ceratium floras
(chart 54). Two of these regions are in the North Atlan-
tic, three are in the Pacific. They are also character-
ized by particular hydrographlc conditions.

Region 1. Cold North Atlantic (stations 3 to 13). --
This region Includes the loop of eleven stations lying
north of 40° north and extending to Iceland. The entire
area is composed of eutrophic water with surface tem-
peratures below 16° C and the phosphate content of the
upper 50 meters above 20 mg P04/m3. The Ceratium
flora in this region is characterized by subpolar species
as well as by tolerant tropical species which probably
have been carried northward by the West Wind Drift and
its tributaries. It is also characterized by a paucity of
species (see p. 6), although the total population is rela-
tively high. The number of species at each station was;
ten at two stations, less than this at the rest, and as low
as one and two at some.^ The total number of species
found In the region was only fourteen.

The following species were found at five or more of
the eleven stations in the region. They are either sub-
polar or cosmopolitan species.

C. arcticum
v. arcticum at six stations
V. longipes at seven stations
v. ventricosum at five stations

C. furca

C. fusus

C. horridum v. horridum

C. lineatum

C. macroceros subsp. macroceros

C. tripos subsp. atlanticum

The following species were found at only one to
three stations in the region. They are all tolerant tropi-
cal or cosmopolitan species.

C. arietinum subsp. bucephalum

C. azoricum

C. declinatum

C. extensum

C. hexacanthxun

C. horridum var. molle

C. macroceros subsp. gallicum

C. massiliense

C. ranipes

Thus, the dominant flora of region I is subpolar in
character and a subdominant element is made up of tol-
erant tropical species. No strictly tropical forms were
found here. The least tolerant forms were C. declina -
tum . C. macroceros subsp. gallicum . and C. ranipes .

'^ to order to make the records comparable, only
oceanc-jraphlc stations are considered in the discussion
of nimibers of species at stations. The records of the
Interposed surface plankton stations have not been used.

CERATroM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

and each of these were found at only one station in the
southern limit of the region.

Region 11. Warm Atlantic (stations 1 to 2. 11 to

M-

— This region includes twenty-three stations, all of
which lie south of 40° north latitude. The entire area is
composed of water with surface temperatures above 20°
C. The phosphate content is extremely low everywhere
except at the three northernmost stations where the
mean values for the upper 50 meters were from 14 to
52 mg POl/m^. At the rest of the stations the values
were 10 mg or less. Consequently, except to the north,
the region supports a very small quantity of plankton and
is distinctly oligotrophic. The sparse plankton, however,
is composed of a large number of species. This number
was between ten and twenty per station in the Caribbean
Sea and western part of the North Equatorial Current,
but higher at the other stations, being between twenty
and thirty species per station at all stations but station
16 where there were thirty-three. The total number of
species found in this region was forty -six.

The Ceratium flora in region n is distinctly tropi-
cal, with the usual number of cosmopolitan forms and a
few subpolar occasionals. The following species were
found at fifteen or more of the twenty-three stations.
This lists represents only intolerant and slightly toler-
ant forms except for two very tolerant species (C. hexa-
canthum and C. massiliense) and two cosmopolitan spe-
cies (C. macroceros and C. tripos) .

C. breve
C. candelabrum
C. contortum
C. contrarium
C. euarcuatum
C. gibber um
C. gravidum
C. hexacanthum
C. macroceros

subsp. macroceros at three stations

subsp. gallicum at twenty-one stations
C. massiliense
C. ranipes
C. tenue

V. inclinatum at seventeen stations

V. tenuissimum at seven stations
C. tripos

subsp. atlanticum at eight stations

subsp. semipulchellum at nineteen stations
C. vultur

V. sumatranum at fifteen stations

V. pavillardii at eight stations

V. japonicum at eight stations

V. vultur at eleven stations

V. recurvum at ten stations

The following species were found at from five to
fourteen stations. They consist of tropical and cosmo-
politan species as well as one subpolar.

C. arcticum

V. arcticum at six stations

V. longirostrum at six stations

V. ventricossum at one station
C. arietinum

subsp. arietinum at four stations

subsp. gracilentum at two stations
C. carriense
C. declinatum
C. extensum
C. furca
C. fusus

C. horridum

V. horridum at five stations

v. molle at nine stations

V. claviger at one station
C. limulus
C. lunula
C. paradoxides
C. pulchelltun
C. reflexum
C. symmetricum

V. symmetricum at one station

V. coarctatum at nine stations

V. orthoceros at four stations
C. teres
C. trichoceros

The following spec: s were found at less than five
stations in the region. This list comprises intolerant
tropical species almost exclusively.

C. azoricum C. belone

C. cephalotum C. concilians

C. digitatum C. falcatum

C. humile C. incisum

C inf latum C. lineatvim

C. longirostrimi C. longissimum
C. subrobustum

The relative frequency of the species in this region
is approximately the same as that for the collection as &
whole so that no particular significance can be attached
to it. The species that are rare in region n are rare
throughout the world and vice versa.

The southern part of region n corresponds to the
northern part of Peters' (1934) region I of the South At-
lantic. The composition of dominant forms in the Car -
negie material agrees with that of Peters.

The area with a low n\miber of species per station
from stations 26 to 34, including the Caribbean Sea, can-
not be considered biologically different except In its
greater general poverty. The temperatures are higher
there, all above 27° or 28° C, and the plankton generally
sparse. Although the number of species per station is
generally low throughout this area, the total number for
the area is not, on the basis of number of stations in-
volved, much different from the total for region I. The
species not represented are mostly the rare forms.
Thus it is probably simply a case of a very poor produc-
tion of all species so that the rarer forms are seldom
collected.

Region m. Cold North Pacific (stations 116 to 128) .
--This region includes thirteen stations, all but one of
which lie north of 40° north latitude. The area is com-
posed of cold eutrophic water with surface temperatures
below 17° C (as low as 7.°15 at one station), and the phos-
phate content of the upper 50 meters above 25 mg
P04/m3 at all stations but one and above 100 mg at the
seven northernmost stations. Consequently this region
supports a dense growth of plankton although the number
of species is small. This number is below ten at all but
one station.

The Ceratium flora in this region is characterized
in general by a few subpolar and cosmopolitan species
and at the southernmost stations there are a few records
of slightly tolerant and very tolerant tropical species.

The following species were foxmd at more than three
of the stations. They are either subpolar or cosmopoli-
tan species.

C. arcticum
V. arcticum at two stations
V. longlpes at nine stations

HORIZONTAL DISTRIBUTION OF CERATIUM

C. arietinum
subsp. arietinum at six stations
subsp. bucephalum at two stations

C. fusus

C. pentagonvun
subsp. pacificum

C. tripos
subsp. atlanticum at seven stations
subsp. semipulchellum at one station

The following species were found at three or less
stations in the region. This list is composed of subpo-
lar, cosmopolitan, very tolerant, and slightly tolerant
tropical species.

C. azoricum
C. carriense
C. contrarium
C. furca
C. horridum

V. horridvun
C. massiliense
C. tenue

V. inclinat\un

C. candelabrtmi
C. concilians
C. extensum
C. gravidum
C. lineatum
C. macroceros

subsp. gallicum
C. petersii

Region IV. Warm Pacific (stations 45 to 47. 63 to
66. 78 to 115. and 129 to 160) . --This region includes
eighty-seven stations covering the tropical Pacific and
extending to approximately 40° north and 35° south lati-
tudes except in the southeast Pacific where it meets the
southeast Pacific region. Region IV is characterized by
warm water with surface temperatures above 20° except
at the borders of the region where it is lower, particu-
larly off California where stations with surface tempera-
tures as low as 16.°3 have been recorded. The phosphate
content of the water in this region Is not the same
throughout. The central part of the North pacific is
practically devoid of phosphate and mean values for the
upper 50 meters is under 10 mg P04/m3, At the north-
em limits of the region these values increase to 25 mg.
The phosphate content of the water in the southern part
of region IV is mostly above 25 mg except in the Easter
Island Eddy where it is between 10 and 25 mg.

Region IV is characterized by a large number of
species of Ceratium per sta^on although the total plank-
ton population is low on the whole. The number of Cera-
tium species per station is everywhere between twenty
and thirty except for four stations near the equator
where it is over thirty and at the northern transitions
where it is between ten and twenty.

The Ceratium flora of this region is distinctly tropi-
cal and includes all the strictly tropical species as well
as the tolerant ones and cosmopolitan and stray subpolar
forms.

The following species were found at more than sixty
stations in this region. This list is composed principal-
ly of slightly tolerant and very tolerant tropical species.

C. candelabrum

C. carriense

C. contortum

C. contrarium

C. declinatum

C. euarcuatum

C. extensum

C. gibberum

C. gravidum

C. hexacanthum

C. macroceros subsp. gallicum

C. massiliense

C. tenue
v. inclinatum at fifty -eight stations
V. tenuissimum at twenty-one stations

C. tripos
subsp. atlanticum at twelve stations
subsp. semipulchellum at seventy -five stations

The following species were found at twenty-five to
sixty stations in the region. This list is composedmost-
ly of Intolerant and slightly tolerant tropical species.

C. arietinum

subsp. arietinum at sixteen stations

subsp. bucephalum at fifteen stations
C. bigelowii
C. breve
C. cephalotum
C. concilians
C. deflexvun
C. falcatum
C. furca
C. fusus
C. limulus
C. lunula
C. paradoxides
C. pentagonum

subsp. tenerum
C. praelongum v

C. platycorne
C. pulchellum
C. ranipes
C. subrobustimi
C. symmetricum
C. teres
C. vultur

v. sumatranum at thirty-four stations

V. regulare at one station

V. reversum at six stations

V. pavillardll at seven stations

V. vultur at sixteen stations

V. japonicum at eighteen stations

The following species were found at less than twenty-
five of the stations in the region. This List, also, is com-
posed principally of Intolerant and slightly tolerant spe-
cies.

C. arcticum v. longlpes

C. aultii

C. axlale

C. azoricum

C. belone

C. bohmii

C. carneglei

C. digltatum

C. filicorne

C. genlculatum

C. horridum

V. horridum at ten stations

v. claviger at ten stations

V. moUe at seven stations
C. incisiun
C. inf latum
C. kofoidii
C. longirostrum
C. petersii
C. reflexum
C. setaceum

Reeflon V. Southeast Pacific (stations 35, 36. 38 to
44. 58 to 62. and 67 to 77) .--This region includes twenty-
five stations in the southeastern Pacific from Panama to
40° south latitude and from the South American coast

6

CERATIUM m THE PACIFIC AND NORTH ATLANTIC OCEANS

westward. It has two tongues extending westward, one
to the north, and one to the south of the Easter Island
Eddy. Although the temperatures vary a great deal in
this area, the water is everywhere eutrophic. The sur-
face temperatures vary from 14.°97 C at the southern-
most station to 27.°4 in the Panama region. The mean
phosphate content of the upper 50 meters is everywhere
above 25 mg P04/m3 and is above 50 mg at all stations
except those bordering the Easter Island Eddy.

The waters of this region support a large plankton
population and, characteristically, few species are pres-
ent. Region V has been delimited by the points at which
the number of species per station rises above twenty.
Not only is the number of species lower in this region
than in the warmer tropical waters to the west, but the
composition of the Ceratixim flora is somewhat different.
The following tropical species found to the west were ab-
sent from region V. They are, significantly, all intoler-
ant tropical species.

C. axiale
C. digitatimi
C. inf latum
C. reflexum

C. cephalotum

C. filicorne

C. longirostrum

The following species were found at fifteen or more
stations within the region. This list is composed mostly
of cosmopolitan and tolerant tropical species.

C. candelabrum
C. contortum
C. furca
C. fusus
C. horridum

v. horridum at twelve stations

V. molle at five stations

V. claviger at four stations
C. lunula
C. massiliense
C. pentagonum

subsp. tenerum
C. tripos

subsp. atlanticum at ten stations

subsp. semipulchellum at fifteen stations

The following species were found at five to fourteen
stations in the region. They are mostly slightly tolerant
tropical species.

C. arietinum
subsp. arietinum at nine stations
subsp. gracilentum at three stations
subsp. bucephalum at three stations

C. azoricum

C. breve

C. carriense

C. concilians

C. declinatum

C. deflexum

C. extensum

C. gibberum

C. gravidum

C. hexacanthum

C. llmulus

C. macroceros subsp. gallicum

C. ranipes

C. symmetrlcum

C. tenue
V. Inclinatum at four stations
V. tenuisslmum at two stations

C. teres

C. trichoceros

C. vultur

V. vultur at two stations

V. pavillardii at two stations

V. reversum at seven stations

v. regular e at one station

The following species were found at less than five
stations. They are notably almost all intolerant tropi-
cal species.

C. belone
C. carnegiei
C. euarcuatum
C. incisum
C. longissimum
C. petersii
C. praelongumi
C. setaceum

C. bohmii
C. compressum
C. falcatum
C. kofoidii
C. paradoxides
C. platycorne
C. pulchellum
C. subrobustum

It is evident from the above lists that the most fre-
quent forms in region V are either cosmopolitan or tol-
erant tropical species. The intolerant tropical forms
are rarely collected in the region so they compose the
bulk of the list of rare forms.

Richness of Species in Different Areas

The well-known difference in number of species
found in cold and warm areas of the oceans is well illus-
trated by the Carnegie Ceratium data (see chart 53). In
the cold North Atlantic the number of species found at
each station was everywhere less than ten. This area
resembles the floristic region I, cold Atlantic.

South of this area there is a transition zone in which
the number of species per station is between ten and
twenty. This lies in region II, the warm Atlantic region.
The rest of this region is characterized by high number
of species, more than twenty, except in the western part
of the North Equatorial Current, from longitude 40 ° west
to Panama, where the values were equal to that of the
transition zone, namely, between ten and twenty species
per station.

In the Pacific the cold northern water is character-
ized by less than ten species per station. This area is
almost exactly like the floristic region IE, cold North
Pacific. At the southern limits of this area in both the
west and east there is a transition zone with values be-
tween ten and twenty species per station.

These transitions lie within region IV, warm Pacific.
The rest of this region has more than twenty species per
station and, at some stations, more than thirty.

In region V, southeast Pacific, the values are mostly
transitional in amount, namely, between ten and twenty
species per station. At one station west of the Panama
area (station 37) there were more than twenty species.
It is possible that this station has not been rightfully in-
cluded in region V but belongs to region IV, the warm
Pacific. The surface temperature at this station was
higher than at any other station in region V, namely,
27.°12 C. The area at this latitude between this station
and the central Pacific is unknown oceanographlcally and
planktologically so that it is not possible to say whether
station 37 lies in an eastern extension of region IV or
not.

South of this station there were two stations at
which the number of species per station was less than
ten, stations 38 and 39. These were the only stations in

FACTORS AFFECTING HORIZONTAL DISTRIBUTION

latitudes lower than 40° where the number of species
per station dropped below ten.

The relation between the richness of species and

environmental conditions will be taken up in the follow-
ing sections.

FACTORS AFFECTING THE HORIZONTAL DISTRIBUTION OF CERATIUM

Salinity

It is probable that the slight variations in salinity
found in the oceanic waters have no influence on the dis-
tribution of Ceratium species. Peters (1934) coulddem-
onstrate no effect in the South Atlantic nor could Nielsen
(1934) in the South Pacific. Nielsen did find that in the
Panama region, where the salinity is low, the total num-
ber of species was low. Conditions were neritic there,
however, so it was impossible to determine which was
the influencing factor. In the Carnegie investigations no
correlation could be found between salinity and the dis-
tribution of any species. In this connection it is worthy
of note that the boundary between the distribution of the
warm and cold water species of the North Atlantic is ap-
proximately at the 36 per mille isohaline; in the North
Pacific the limits of the same species occur at 34 per
mille off Japan and 33 per mille off California. Thus,
apparently these species are not affected by such slight
variations in the salt content as are found In the open
ocean.

Nutrient Salts

Since the depletion of phosphate and nitrate is known
to limit the total production of phytoplankton in some lo-
calities, it is important to examine these factors in rela-
tion to the world distribution of Ceratium species. Ni-
trate determinations were not made on the Carnegie but,
since the fluctuations in nitrate and phosphate run in a
parallel manner, it is sufficient to discuss the phosphate
distribution as representative of the inorganic nutrient
materials.

In general, regions of very low phosphate content
are characterized by sparse populations of all species
but by a larger number of species. This led Peters
(1934) to suggest that nitrate and phosphate inhibited the
growth of certain species. Thus in region I of the North
Atlantic where the phosphate content of the upper 50 me-
ters is above 25 mg P04/m3, the number of Ceratium
species is less than ten per station (see charts 52 and
53). In region n the phosphate content is under 10 mg
and the number of species of Ceratium per station every-
where above ten and, usually above twenty, and at one
station above thirty. Likewise in the Pacific the phos-
phate-rich regions are characterized by a poor Ceratium
flora. Region HI, where the concentrations run above 25
mg and at most stations above 100 mg, the number of
species per station is less than ten. The correlation is
not quite so good In region IV, the warm Pacific region.
The number of species per station is between twenty and
thirty throughout most of this region although the phos-
phate content of the northern part is under 10 mg and in
the southern part it varies from 10 to 50 mg. The south-
east Pacific region has many values above 50 mg, that
Is, higher than the warmer region to the west. Accord-
ingly the number of species per station is less, namely,
between eleven and twenty.

Thus, within any given area there seems to be a
correlation between high phosphate content and small
number of species. It is important to note, however,
that the limit between any particular number of species
does not occur at the same concentrations of phosphate.
Thus, in the northern Atlantic less than ten species are
found in water with 25 to 50 mg of phosphate, whereas
in the South Pacific the area with more than twenty spe-
cies includes water with over 25 mg P04/m3. Similar-
ly, in the North Pacific all the water containing more
than 50mgPO4/m3 has less than ten species per station,
whereas in the southeastern Pacific there are areas
where the phosphate content is above 50 and the species
per station are between ten and twenty.

The same sort of correlations can be found between
the Ceratiimi life zones and phosphate content. Each re-
gion has its characteristic phosphate content. Thus re-
gions I, ni, and V have high concentrations, whereas
region II has uniformly low concentrations, and region
IV is low in the north and has medium values in the
south. As in the case of number of species, the correla-
tion with floras is not exact. Thus the flora of region I
has little in common with the flora of the southern part
of region IV which has the same phosphate content.

These studies tend to indicate that the phosphate
content of the water has no direct effect on the horizon-
tal distribution of Ceratium species, at least not as re-
gards absolute values. There are indications, however,
that the relative values in a given region bear some re-
lation to the Ceratiimi flora. Perhaps some factor as-
sociated with an increase in phosphate is significant in
the distribution of Ceratium species.

Temperature

Peters (1934) came to the conclusion that water
temperatures between 15° and 27.°5 C had no influence
on the distribution of Ceratia in the South Atlantic. How-
ever, he was considering the yearly range of tempera-
ture at each locality. Peters was able, on the other hand,
to group his species according to thermal environments.
Of his fifty-five species, thirty-three were limited to
warm water, twenty-one to warm and cool water, and
only one was observed in the southern cold water. Niel-
sen (1934) was inclined to consider 15° C too low for
noneffective temperatures. In the. regions which he in-
vestigated in the South Pacific, however, he did not find,
many cases of temperature correlation. He designated
only two temperate forms-- C. petersii and C. tripos at-
lanticum. He also found that C. filicorne is restricted
to high temperatures in the South Equatorial Current.
Nielsen, however, did classify his species according to
temperature zones with the following categories: tropi-
cal, tropical-subtropical, tropical-subtropical-temperate,
and temperate.

In the Carnegie collections it is possible to study
the transition from tropical to cold water regions infour
different places: In the western North Atlantic, western

8

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

North Pacific, eastern North Pacific, and eastern South
Pacific. A comparison of the Ceratia distributions in
these regions Is very Interesting, and gives us a clue to
the factors controlling distribution, at least in the case
of some species. For these studies it is important to
consider first the most common tropical species so that
negative records in the transition zones can be given
some weight. Ceratium contortum is a good example of
a tropical distribution (chart 32). The species drops out
at surface temperatures of about 20° in the North Atlan-
tic, at above 20° off Japan, 15° off California, and 15° in
the southeastern Pacific. Many other species have a
similar distribution, some more closely confined to cer-
tain temperatures and some less. Ceratium massillense
(chart 38) is a common tropical form whose limits of
distribution parallel rather closely the 15° C isotherms
in the four transitions. Some species show a definite
restriction to tropical water except for a particular dis-
placement by a current. A good example of this is Cera-
tium hexacanthum which is within the 20° isotherm at
the three transitions in the Pacific, but In the Atlantic it
ts apparently carried by the currents to Iceland where
the surface temperature is less than 10°. Conversely,
there are species which are restricted to warm water
throughout their ranges. Examples of these are C. breve
(chart 21) and C. lunula (chart 31).

A few forms are found in the cold water but not in
the tropics, or only in insignificant numbers there.
Ceratimn arcticum (chart 47) is an example. Still a
third general type of distribution Is found in such forms
as C. fusus (chiairt 17) which occur In cold water as well
as in warm, and are truly cosmopolitan.

On the basis of the distributions of the Ceratium
species at the Carnegie stations, a classification con-
sisting of three main categories was devised: tropical,
subpolar, and cosmopolitan. The tropical, in turn, was
divided into three regions. Those species which were
rather closely restricted to surface temperatures of 19°
or above were grouped together as intolerant tropical
species. Those which transgressed a little into the
transition zones were called slightly tolerant tropical
(orms and, finally, those which were carried far beyond
the tropical regions were designated very tolerant tropi-
cal forms. This classification has no category for tem-
perate species such as most previous classifications
had. The authors believe that there are no truly temper-
ate oceanic species of Ceratium. The temperate lati-
tudes are populated by tropical and cosmopolitan forms
with occasional appearances of subpolar forms. Follow-
ing is the geographic classification of the Carnegie
Ceratia based on the above scheme.

Intolerant

Tropical

c

inclsum

C. longirostrum

c.

inflatum

C. falcatum

c.

praelongum

C. cephalotum

c.

dlgitatum

C. belone

c.

blgelowii

C. breve

c.

euarcuatum

C. limulus

c.

paradoxides

C. vultur

c.

longissimum

C. reflexum

c.

deflexum

C. lunula

c.

fllicorne

C. trlchoceros

c.

axlale

C. genlculatum

c.

hum lie

Slightly Tolerant Tropical

c.

subrobustum

C.

teres

c.

kofoidii

C

gravidum

c.

pulchellum

c.

symmetricum

c.

declinatum

c.

gibberum

c.

concilians

c.

ranipes.

c.

carriense

c.

platycorne

c.

tenue

c.

contrarium

c.

macroceros subsp.

c.

pentagonum subsp

gallicum

tenerum

c.

candelabrum

c.

contortum

c.

tripos subsp.

c.

aultli

semipulchellum

c.

setaceum

c.

carnegiei

c.

bohmil

Very Tolerant Tropical

C. azoricum C. massiliense

C. arletinum C. extensum

C. hexacanthum

Cosmopolitan

C. horrldum C. fusus

C. furca C. petersii (?)

C. tripos subsp. atlantlcum

Subpolar

C. llneatum C. macroceros subsp.
C. pentagonum subsp. macroceros

pacificum C. articum
C. compressum (?)

Currents

Currents are certainly important in the distribution
of Ceratia. They cause displacements in the normal
distribution by carrying populations into regions where
they have not developed, and they create displacemtBts
of the normal range of environmental conditions so that
species can develop in regions which otherwise would
not be favorable for them. The current factor in distri-
bution, however, is the most difficult one to demonstrate
by a gross world survey such as that of the Carnegie .
The value of Ceratia as current indicators is undoubted-
ly much greater than the Carnegie data would indicate.
The importance of such indicators is evidenced most in
temporary current anomalies which can be observed
satisfactorily only l>y a continued study of a particular
area.

It shoxild be emphasized that a current which dis-
places a flora from its normal 'distribution also displaces
the environmental conditions. When the displaced water
mixes with the adjacent water, it may no longer be suit-
able for the organisms displaced and they disappear.
That the amoimt of mixing in such currents is great is
indicated by the rather sharp delimitation of many of the
Ceratium species in the Carnegie collection. This is
particularly striking in the Kuroshio and Gulf Stream,
two currents well known for their velocity and general
Influence. In these currents the tropical floras change
their fades rather rapidly when the surface tempera-
tures drop below 19° C.

Peters (1934) considered that ocean currents played
a great role in the distribution of Ceratia in the South
Atlantic. He was able to give comparatively few in-
stances of displacements of species, however, in the
areas he studied.

FACTORS AFFECTING HORIZONTAL DISTRIBUTION

9

The authors tend to agree with Gran (1912) that the
nature of a Ceratlum flora is determined more by the
chemical-physical conditions of the water than by trans-
portation by currents. The fact that a warm current
carries a tropical flora into high latitudes is not opposed
to this argument, as present evidence indicates that such
a current will do this principally by carrying the tropi-
cal environment with it.

In the Carnegie investigations there were four re-
gions where current influences would be expected to be
demonstrated. These concerned four great currents:
the Gulf Stream, Kuroshio, California, and Humboldt
currents.

In the North Atlantic West Wind Drift or "Gulf
Stream" the intolerant tropical species dropped out at
about 45° north. The best examples of displacement,
however, were found here in the case of some of the tol-
erant tropical species. Thus, C. extensum was carried
to the British Isles to temperatures of 12.° 44 C (chart
16) and C. hexacanthum to Iceland in temperatures of
8.°92 C (chart 48). Unquestionably these are examples
of current displacements as C. extensum was not found
elsewhere in temperatures below 14.°97 nor C. hexacan-
thum below 18.°97. The occurrence of C. massiliense and
C.platycorne off Ireland should probably also be consid-
ered displacements (charts 38 and 35). Other tropical spe-
cies such as C. declinatum and C. ranipes were carried
over into region I only as far as station 3, where the
surface temperature was 15.°5 C (charts 28 and 36).

In the corresponding current of the Pacific, the
Kuroshio, we find that the transition from tropical to
subpolar floras is more abrupt. Most of the tropical
species were not carried into water with temperatures
under 20 ° C . Only C. gravidum . C. candelabrum , and C.
tripos semipulchellum . of the tropical species, were
carried into water with surface temperatures as low as
15.°93 C (charts 3, 5, and 19). No evidence of any tropi-
cal species being carried into lower temperatures than
that in this region was found.

Corroboration of the lack of displacement in the
Kuroshio was found in the distribution of the tropical
species at similar latitudes in the western Pacific off
California. If the currents were effective In displacing
the distributions significantly into different environ-
ments, then the tropical species should be found in cold-
er water off Japan than off California. The Kuroshio
should sweep populations into cold water and the Cali-
fornia Current should push the northern limit southward.
This was seldom found, however. The species which
disappeared at 19° C off Japan, reappeared at 19° C off
California. There were many variations between spe-
cies but, in the main, the tropical flora of the North Pa-
cific had its northern limits in the western and eastern
Pacific at the same isotherm, and this isotherm was at
approximately the same latitude, namely 40° north.

There were two striking exceptions to this general
distribution, however; the distributions of C. breve
and C. lunula (charts 21 and 31). These species were
found rather consistently within tropical latitudes, that
Is, within 20° of either side of the equator. The ranges
of these species beyond these latitudes, however, does
not follow that of other common tropical species which
are usually found throughout the warm-water regions to
about 40° north and south. In the case of C. breve the

records do not extend far north or east of Hawaii,
although in the western Pacific they extend to Japan.
They are absent from a large part of the southeast Pa-
cific region, although they extend to Easter Island. The
distribution of C. lunula is stUI more uneven in the
North Pacific. In the east it was found nowhere north of
20°, whereas in the west it was found continuously to
northern Japan, latitude 38° north. In the South Pacific,
however, it was everywhere within 20° south latitude.
Is it possible that these distributions are determined by
the current systems? Certainly such distributions can-
not be accounted for by simple current displacements as
a bodily displacement of the plankton would have dis-
placed other species as well. Could these curious dis-
tributions be the result of a specific susceptibility to
something? Is the absence of these species in the south-
east Pacific owing to some deleterious effect of the out-
wash of subantarctic and upwelled water from the south
and east, and is their absence from the region between
California and Hawaii owing to the combined effects of
the California Current and upwelling? The answers to
these questions cannot be given until further work Is
done. It is remarkable that only two species show such
distributions. The proof of the validity of these must
await further investigations.

The equatorial currents apparently have little effect
on the distribution of Ceratia. This is probably because
they involve the movements of waters of very similar
nature. Although the temperatures of these waters may
vary from 20° to 29° C, the surface temperatures above
20° C apparently have no effect on the Ceratia. Nielsen
found that C. filicorne was found only In the warmest
parts of the South Equatorial Current, but this was not
corroborated by Carnegie data (see p. 28). No segrega-
tion of Ceratlum species within the equatorial regions
could be made on the basis of temperature.

The last great current system to be considered is
the Himiboldt Current off South America. Unfortunately
the Carnegie stations did not run near the continent except
at Callao so the most highly developed part of the current
system was missed. The hydrography of this region is
complicated by a strong upwelling along the coast. The
changes in the environmental conditions brought aboutby
this upwelling are probably the same as those effected
by the Humboldt Current, which brings water from the
subantarctic. These changes consist In lowered temper-
atures and pH, increase in phosphate, etc., and a result-
ant Increase in plankton production. Thus it is impossible
to decide whether the biological conditions peculiar to
this region should be attributed to current phenomena or
to the other hydrographlc feature, namely, upwelling.

Probably there is a displacement by the Humboldt
Current of antarctic forms northward along the southern
part of the South American coast southeast of the Carne-
gie track, but no evidence of this was observed at any
Carnegie station. Probably the only antarctic form of
Ceratlum is C. pentagonum robustum (Peters, 1934).
This did not occur in the Carnegie collection.

The Humboldt Current is deflected westward, and
there is a more or less general movement of water away
from the South American coast in the southeastern Pa-
cific. How far westward the influences of this movement
are felt is an interesting problem. In the discussion of
the hydrographlc conditions, it was pointed out that the

10

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

influence on the chemical and physical conditions of the
water are far-reaching. Associated with this we find
that the biological conditions have been affected for
hundreds of miles offshore. In the first place there is
a high production of total plankton; secondly, there is a
reduced number of species per station in the region
(see chart 53); and thirdly, the development of some of
the tropical species apparently has been suppressed.

Many of the tropical species common in the waters
to the west were absent in at least certain parts of the
southeastern Pacific. Sometimes this distributional hia-
tus was confined to the Peruvian coast and Galapagos
area as in C. extensum (chart 16) and C. tenue (chart
45). Other species seemed to show a more or less gen-
eral avoidance of the area, as for instance, C. pulchel-
lum (chart 20), C. carriense (chart 40), C. vultur (chart
43), and C. teres (chart 11).

Here again we probably are not confronted with a
problem of simple current displacement, but rather with
current effects brought about through a modification of
the environment. What these peculiar modifications
might be, will be discussed in the next section.

Other Factors

Although the above geographic classification of the
Ceratia (p. 8) has a temperature connotation, it is not
intended to mean that the controlling factor in Ceratium
distribution is necessarily temperature. There are stlU
other factors which are associated with temperature
which we have not yet discussed.

Nielsen (1934) is of the opinion that the concentra-
tion of plankton organic metabolic products is the most
important factor in the distribution of Ceratium species
in the warmer seas. Since no measurement of such
substances are available, it is impossible to test this
theory. Nielsen based his opinion principally on the
distribution of neritic and oceanic species and on the
relative number of species in these two environments.
He emphasized the fact that In neritic waters only a
small number of Ceratia occur. That this is not owing
to higher concentrations of nutrients alone was shown by
the Great Barrier Reef region where the nutrients are
low as well as the Ceratium flora. Nielsen found higher
concentrations of nutrients in the open ocean (eastern
South Equatorial Current) and a diminution in species.
He contended that the metabolic products require con-
siderable time for decomposition, that the fertility of

neritic water is replenished from shallow depths where
these products have accumulated, and that in oceanic re-
gions no such accumulation can occur. What these met-
abolic products are, remains to be demonstrated.

Even If the presence of such products is demon-
strated and correlations found, their causative nature
must be proved and this probably will have to be done
experimentally.

The difficulties In establishing the important envi-
ronmental conditions are accentuated by the fact that
many water conditions change together. The agency
which enriches the photic zone to any great degree is a
movement of water from deeper levels to the surface
layers. This one agency, whether near shore or in the
open ocean, brings about a whole set of new environ-
mental conditions. Not only is the phosphate and nitrate
content of the water Increased, but the temperature is
lowered and the hydrogen-ion concentration Is increased.
Undoubtedly other changes are effected also. If some
particular metabolic product Is rather stable, as Nielsen
postulated, no doubt higher concentrations of this accom-
pany any enrichment of the photic zone. A point worth
considering in this connection, however, is thatNlelsen's
postulated metabolic product poisoning may be purely of
plant origin. In this connection Hardy's (1935) discus-
sion of the "exclusion effect" of phytoplankton for zoo-
plankton is interesting. Considerable evidence is being
acciunulated that Indicates that phytoplankton exerts an
exclusion effect on many animals. Is it possible that, in
a similar way, it may exclude certain oceanic species of
Ceratium?

The association of all these factors makes it diffi-
cult to select the particular one which might be influenc-
ing the dlstrlbutlpn of any particular species. Thus the
floristlc zones outlined above are separated not only by
certain temperature differences, but also by certain as-
sociated differences in phosphate, pH, plankton content,
etc.

The difficulty Involved In determining causative fac-
tors is well illustrated by the waters of the southeast
Pacific where, obviously, some condition associated with
the Humboldt Current and upwelllng limits the distribu-
tion of certain Ceratium species. This condition here is
probably not temperature, as many of the temperature
records were over 20°. Which of the conditions associ-
ated with cold currents and upwelllng is the determining
one. however, cannot be pointed out at this time.

COMPARISON OF THE ATLANTIC AND PACIFIC WATERS
ON THE BASIS OF THEIR CERATIUM FLORAS

There Is no known difference in the environmental
conditions of the waters of the Atlantic and Pacific
oceans which might operate to favor or hinder certain
species in one ocean and not in the other. Thus it would
be expected that, given the same plankton commimlties,
the same species would survive in each. The two oceans
are completely Isolated from each other, however, except
in the south, where they both merge Into the Antarctic
Ocean, and In the far north, where they are connected by
way of the Arctic Ocean and the narrow Bering Strait.
Since temperature or some correlated factor Is probably

responsible for the distrlbirtion af Ceratium species,
temperature will be discussed in connection with the
comparison of the floras of the two oceans. The surface
temperatures In the latitude of Cape Horn In summer
are under 8° C. Whether 8° is sufficiently low to con-
stitute a barrier to tropical Ceratium species or their
spores, is unknown. The continental barrier between
the Atlantic and bdlan oceans certainly must be less
effective since the temperatures at the Cape of Good
Hope In summer may be above 20° C. Thert Is no con-
tinental barrier between the Indian and Pacific oceans.

COMPARISON OF ATLANTIC AND PACIFIC WATERS

11

It would be supposed that any species originating in the
Atlantic would, in time, be swept to the Indian and Pa-
cific oceans since the currents in the far southern
waters are predominantly from west to east. Whether
the Pacific tropical species could find their way to the
Atlantic by the southern connection is less certain.

The water connection of the two oceans to the north
by way of Bering Strait and the Arctic Ocean certainly
would be expected to constitute a barrier to Ceratium
species, at least to the tropical forms. Bering Strait
is shallow and narrow, with water temperatures around
8° C. It is not likely, however, than an organism could
pass from here to the Atlantic without passing through
water of 0° C or lower.

In the light of this knowledge it is interesting to
compare the Ceratium floras of the Pacific and Atlantic
oceans. An examination of the Carnegie lists of species
for the two oceans shows that most of the species are
common to both oceans. Nevertheless, there are some
significant differences. Nine of the species occurred in
only one ocean and this, significantly was the Pacific.
One of these, C. axlale . has been foimd in the Atlantic by
others, so it will be disregarded. The others are as fol-
lows: C. aultii. C. bigelowli . C. carnegiei . C. deflexum.
C. fllicorne . C. geniculatum . C. peter sii. C. bohmii.

None of the Atlantic species were absent from the
Pacific. In Nielsen's comparison of the two oceans, he
cites two examples of Atlantic forms which are absent
from the Pacific, namely, C. longlnum Karsten (=C.
arcuatum longlnum In Peters, 1934), and C. mlnutvun
Jorgensen. These forms, however, have now been found
abundantly in the Pacific in the Carnegie collections (see
pp. 35 and 22). Otherwise Nielsen's comparison of the
two oceans agrees well with our own, except that he lists
C. himiile as absent from the Atlantic. It was found in
that ocean in the Carnegie collection.

K is true, of course, that the species peculiar to the
Pacific are rare forms and may yet be discovered in the
Atlantic. In the case of at least three of the species,
(C. bigelowli. C. deflexum . and C. fllicorne) . however,
this is not likely, as many records of their occurrence
are now being acctmiulated. For Instance, in the Carne-
gie collection alone there are thirty-seven sample rec-
ords for C. bigelowli . one hundred five for C. deflexum.
and twenty -seven for C. fllicorne.

R shoxild be noted that the eight species peculiar to
the Pacific are strictly, or only slightly tolerant, tropi-
cal species, not occurring at stations where the surface
temperature was less than 20° C. Thus, these species
might well find it impossible to pass around Cape Horn
in the cold southern water.

Therefore, it seems quite possible that the above-
mentioned seven species have originated in the Pacific
Ocean and must forever remain in the warm regions of
that great ocean, being barred from the Atlantic by the
great southern extension of South America, whereas the
forms that have originated in the Atlantic have found an
easy migration to the Pacific by way of the Indian Ocean.

We must also bear in mind that the other species of
the two oceans may not be as similar as present work-
ers believe. After all, the morphology of Ceratium is
known only grossly; very few species have been anal3^ed
in detail, and none of them completely. Perhaps real
specific characters have been overlooked. In this con-
nection it is important to examine some differences in
the distribution of some of the species which, according
to present taxonomic methods, are specifically Identical

in the two oceans. Five of the tropical species were
decidedly more tolerant to cold water in the Atlantic
than in the Pacific; namely, C. furca. C. extensum . C.
horrldum . C. hexacanthum . and C. tripos anan^ j niir ,
whereas one cold water species, C. arcticum was appar-
ently more tolerant to warm water in the Atlantic than
in the Pacific (see charts 6, 16, 44, 48, 18, and 47).
Ceratium horridtmi . in addition, seemed to show a dif-
ference in its relation to ollgotrophlc water in the two
oceans, being found practically only in eutrophlc water
in the Pacific but in ollgotrophlc as well as eutrophlc
water in the Atlantic. These differences in distribution
in the two oceans strongly suggest that the genetic com-
plex of the forms in the two oceans is not identical. The
representatives of the two oceans may be different spe-
cies or subspecies or maybe "physiological subspecies."
The solutions to these problems must await a more mi-
nute morphological examination of the forms in question.

About 20 per cent of the tropical species of Ceratium
in the Carnegie collection show either morphological or
distributional features peculiar to one ocean, indicating
an isolation of the tropical waters of the Atlantic and Pa-
cific. The evidence for the isolation of the North Pacific
cold-water region, however, is even more convincing.

Although there are certain similarities between the
Ceratium floras of the cold North Pacific and cold North
Atlantic regions, there are, on the other hand, some very
striking differences which can be accounted for only on
the assumption that these two oceans are biologically iso-
lated from each other. The similarities in the two floras
are expressed by the occurrence in both regions of the
two subpolar species C. lineatum and C. arcticum.

The differences in the floras of the two regions are
more striking than the similarities. In contrast with the
tropical floras these differences are not one-sided, that
is, the "mono-oceanic" forms are not all in one ocean
so that, in contrast with the southern oceans, a mutual
isolation is indicated.

The significant forms in this case are not species,
but subspecies. The first of these to consider is C. furca .
This species, as broadly considered, is cosmopolitan. It
was noticed, however, that in the Atlantic there was a
hiatus between the tropical records and the cold-water
records (chart 6). Although no morphological difference
could be discerned between the southern and northern
forms, it was suggested that they represented at least
physiological subspecies (see p. 18). When we turn to
the Pacific we find that the species is represented only
in the warm-water regions except for one station in re-
g;ion m off Japan, which must be considered a displace-
ment by the Kuroshlo. The species is absent along all the
rest of the cold-water area traversed by the Carnegie .

The case of another species, C. macroceros . is
somewhat more convincing. Inasmuch as the subspecies
are morphologically easily distinguishable. Subspecies
g alUcum is characteristic of all warm-water regions. It
is widespread over both the Atlantic and Pacific. The
species is represented in the cold North Atlantic by sub-
sp. macroceros. In the cold North Pacific, however, the
species is conspicuously absent (see chart 37). An as-
sumption of the isolation of the North Pacific based on
the above two examples alone would not be very conclu-
sive since It would be based on negative evidence In a
region not well Investigated, namely, the North Pacific.
The following example, however, doesnot carry this weak-
ness and, taken with the above two, forms a convincing
proof of the isolation of that ocean.

12

CERATIUM m THE PACIFIC AND NORTH ATLANTIC OCEANS

This example concerns C. pentagomun . a widespread
tropical species. Peters (1934) found a cold-water sub-
species in the subantarctic waters of the South Atlantic,
subsp. robustum . the most southern representative of the
genus. He emphasized the remarkable absence of any
representative of the species in the cold northern waters
of the North Atlantic. The absence of the species in
these waters is well established as the region has been
thoroughly investigated. The Carnegie investigations in
the North Pacific, however, revealed a very divergent
subspecies, subsp. pacificum . which was found only in
the cold North Pacific region (see p. 20). It attained a
high degree of morphological distinctness in the coldest
water, although it intergraded with var. tenerum at its
southern limits. There can be no doubt that subsp.

pacificum is not found in the Atlantic and, therefore, is
peculiar to the cold waters of the North Pacific.

These three striking differences in the Ceratium
floras of the cold regions of the two oceans indicate that
a barrier to at least some of the subpolar species exists
between these two water masses. The nature of this
barrier, of course, is only a matter for speculation.
The hydrography of Bering Strait is very poorly known.
Any continuous current in either direction, however,
would not permit the development of the floras now ob-
taining in the two oceans, providing the species could
traverse the Arctic Sea. Whether they can cross this
ocean is also unknown. An investigation of the phyto-
plankton of the Arctic should throw some light on this
problem.

THE VERTICAL DISTRIBUTION OF CERATIUM

Karsten (1907) first described a special "shade
flora" of the ocean. According to him this consists
principally of Coscinodiscus, Planctoniella, and Goss-
leriella. He thought, however, that the genus Ceratium
was represented at different levels by different species
or varieties and designated C. gravidum Gourret; C^
tripos azoricum CI. var. breve Ostf.. and C. tripos
glbberum Gourret as shade species. As Nielsen (1934)
states, an inspection of Karsten' s lists does not corrob-
orate this idea except In the case of C. gravidum .

Schroder (1911) listed C. inflexum f. claviceps (=C^
contrarium f . claviceps) . C. platvcorne . and C. limulus
as species which "avoid" the intense continued sunlight
of the "southern waters."

Jorgensen (1920) made an Intensive study of the ver-
tical distribution of Ceratium in the Mediterranean. He
found that a great number of species which occurred at
the surface in the winter, inhabited the deeper levels in
the summer. He concluded that the surface summer flo-
ra is more or less indigenous to the Mediterranean,
whereas the winter species are dependent on a migration
from the Atlantic.

Paulsen (1930) and Nielsen (1934) challenged this
migration theory of Jorgensen, but Bohm (1931) accepted
it. Peters (1934) could not establish any vertical distri-
bution from the material of the "Meteor" expedition.

The Dana collected with closing nets at the following
steps: 200 to 100, 100 to 50, and 50 to meters. From
such material it was possible to test the Idea of the
"layering" of the species of Ceratium. Nielsen (1934),
who studied this collection, found that about one -third of
the Ceratium species occurring in the southern Pacific
Ocean must be designated "shade forms," whereas the
rest of the species Inhabit predominantly the upper lay-
ers. He found that the density of the plankton affects the
vertical distribution of the shade forms. They occur in
higher levels in the richer water, presumably because in
such regions there is not sufficient light for growth at
the lower levels.

Nielsen (1934) compared the shade forms of Cera-
tium with the shade plants of the tropical rain forest, all
of which have their leaf surface increased in some way.
The leaves are thin and there is an Increase in assimi-
lating cells. So in Ceratium the cells are thin and crowd-
ed with chromatophores. The cell body (and apical horn)
may be expanded and crowded with chromatophores

(chloraplasts) as in C. gravidum . or the antapicals may
be expanded and rich In chloroplast as in C. platvcorne .
C. claviger . C. ranipes.

Long-horned forms are found among shade forms
as well as among surface forms, but the shade forms
always have the horns crowded with chromatophores.

Of the surface forms there is not a single example
of surface expansion. Thus, the usual assumption, that
surface expansion in Ceratium is a flotation adaptation,
is erroneous.

Since the Carnegie plankton collection contains sam-
ples collected at 50- and 100-meter depths as well as at
the surface, and, since the collection contains practically
all the marine species of the genus, it was possible to
test the theory of Nielsen in the case of each species.
Although the Carnegie collecting nets were open nets,
the duration of towing at the particular level was so much
greater than the time of hauling in, that the percentage
of "contaminants" would necessarily be small. These
contaminants, moreover, would be upper -level forms so
that they would not introduce any error into the calcula-
tions in the case of deep-water forms.

For each species reported in the Carnegie collec-
tion, the number of records for each collecting level was
computed. Since there were more surface hauls than
deeper hauls, these numbers needed to be weighted.
Consequently they were computed as percentages of the
total number of samples collected at the particular depth.
Tables were compiled showing these values, as well as
the actual number of records, for the three levels.
Since an expression of the relative abxmdance of the spe-
cies at each level Is more significant than the mere pos-
itive record, the number of records of "rare," "occa-
sional," etc., and their percentages were computed as
well as the total number, and these were included in the
tables.

Such an analysis of the Carnegie distributional rec-
ords showed that twenty of the species were definitely
more abundant in the deeper levels and nine were ques-
tionably so. All the twenty species showed an increase
In frequency from surface to 100 meters. The agree-
ment between these species and the species indicated as
"shade species" by Nielsen Is great.

There are only three cases of definite disagreement.
Melsen classified C. subrobustum and C. trichoceros as
surface species, whereas the Carnegie data definitely

THE VERTICAL DISTRIBUTION OF CERATIUM

13

Indicate that they are shade species. On the other hand,
Nielsen indicated C. hexacanthum as a shade species,
whereas the Carnegie data failed to substantiate this.
The agreement between the Carnegie and Dana shade
species and the "winter species" of Jorgensen (1920)
is also great. Only one of JOrgensen's winter species,
C. kofoidii . has not been classified as a shade species
by either Nielsen or the present authors. A comparison
of Jorgensen' s "winter species" with the shade species
of Nielsen and the shade species of the Carnegie is given
in table 1.

Nielsen (1934) stated that Jorgensen's (1920) obser-
vations indicated that Ceratia are phototropic; the shade
species in the winter when the light intensity is low
come to the surface and thus are able to maintain a po-
sition in optimal light conditions.

The authors agree that JOrgensen's observations in-
dicated a phototropic response on the part of the "winter
Ceratia," but do not believe that the vertical migration
of Ceratia is a simple light reaction phenomenon. In the
Carnegie collection the shade species were found most
abundantly at 100 meters. Surely the light intensity at
this depth even in summer could not be compared with
the light intensity at the surface in winter in the Medi-
terranean or anywhere else in the world for that matter.
It is scarcely logical to assume on the basis of the pres-
ent data that the shade species of Ceratia seek a zone of
a particular light intensity.

Is it not possible that we have to deal here not only
with a phototropism but a trophotropism as well? In the
summer time the upper levels of the sea are depleted of
nutrient salts to a depth equal at least to that populated
by the phytoplankton. Obviously, the species of phyto-
plankton which are the most tolerant of shade would have
a decided advantage in the quest for nitrogen and phos-
phorus, which occur in the deeper levels in large quanti-
ties. Is it not possible that these forms have a positive
tropic reaction to nutrient salts or some associated con-
dition as well as a phototropic response? By such a
mechanism each species would maintain a position opti-
mal for photosynthesis.

There is another, and simpler, explanation that may
account for the vertical migration of Ceratia. There
may be a simple reversal of phototropic response, de-
pending on the physiological condition of the organism;
in this case on the assimilation of inorganic nutrients or
some associated substance. The mechanism would be
such that with the assimilation of these substances the
organism is positively phototropic, thus remaining near
the surface in the winter; but with the diminution of this
assimilation the organism becomes negatively phototrop-
ic, and thus descends to lower levels of greater nutrient
content.

It must always be borne in mind that the poverty of
nitrogen and phosphorus which land plants everywhere
are fighting, is accentuated to a high degree in the ocean.
V/hen a land plant dies, its nitrogen and phosphorus are
soon returned to the soil to be utilized by other plants.
On the other hand, when a planktonic plant dies, it sinks
below the growth zone and its nutrient elements are lost,
to be returned only after a long period of time except in
high latitudes and in certain peculiar regions. This is
particularly true of the tropics, where the thermal strat-
ification is extreme and continuous. In these regions it
is probable that the fertilization of the photic zone is
accomplished alone by the nocturnal visits of a sparse

zooplankton. E is in such regions as this that the shade
species of Ceratium develop. Nielsen (1934) has already
observed that the shade species are all warm oceanic;
they do not occur in neritic conditions nor in the cold-
water southeast of New Zealand. The Carnegie obser-
vations show that none of the shade species are cold
water species, with the possible exceptions of C. arc-
ticum and C. horrldum . Of the twenty-nine species def-
initely or questionably shade species according to Car-
negie data, seventeen are intolerant tropical species,
eight are slightly tolerant tropical species, only two are
very tolerant tropical species, one is cosmopolitan (C.
horridum) . and one subpolar (C. arcticum) .

The data concerning the depth at which C. arcticum
lives most abundantly are not conclusive, but they sug-
gest that it is a shade species. Ceratium arcticum . a
cold-water species, may be a shade species of another
type entirely. Shiviroff and Federoff (1938) have found
phytoplankton flourishing under the arctic ice cap. They
have not yet reported the species found, but probably C.
arcticiim is one of them as it is characteristic of arctic
currents. In the low illimiination occurring under the ice
it would be expected that only species with a high toler-
ance for shade would be found. The nutrients here are
rich. These forms, when living in exposed ice-free re-
gions, would then have the advantage of the ability to live
at greater depths than the species not tolerant of shade,
so they would be found at depths equal to that of the trop-
ical shade forms.

Table 1. Shade species of Ceratium

Species

In summer,
Mediter-
ranean

Dana
collection

Carnegie
collection

C. praelongum

yes

yes

C. cephalotum

yes

yes

C. gravidum

yes

yes

yes

C, digitatum

yes

?

yes

C. belone

yes

no

?

C. incisum

yes

no

9

C. subrobustum

no

yes

C. kofoidii

yes

no

no

C. setaceum

yes

9

?

C. geniculatimi

•?

7

?

C. bigelowii

....

?

7

C. euarcuatum

yes

yes

yes

C. filicorne

yes

yes

C. symmetricum

yes

yes

yes

C. axiale

yes

yes

C. aultii

?

C. azoricimi

no

?

C. arietinum

yes

yes

yes

C. lunula

yes

yes

yes

C. carnegiei

?

C. paradoxides

yes

yes

C. platycorne

yes

yes

yes

C. ranipes

yes

yes

yes

C. trichoceros

yes

no

yes

C. vultur

a

yes

yes

C. horridum

....

____b

yes

C. tenue

yes

yes

yes

C arcticum

?

C. longissimum

yes

yes

yes

C. hexacanthum

yes

no

C reflexum

yes

yes

? indicates either insufficient data or that the results
were not conclusive. ^Favillardii. ''Molle andclaviger.

14

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS
THE CERATIUM SPECIES OF THE CARNEGIE COLLECTION

Systematic List

I. Subgenus POROCERATIUM

1. C. praelongum (Lemmermann) Kofoid

2. C. cephalotum (Lemmermann) Jorgensen

3. C. gravidum Gourret

II. Subgenus BICERATIUM

4. C. digitatum Schutt

5. C. candelabrum (Ehrenberg) Stein

6. C. furca (Ehrenberg) Dujardin

7. C belone Cleve

8. C. incisum (Karsten) Jorgensen

9. C. pentagonum Gourret

subsp. tenerum Jorgensen
subsp. pacificum n. subsp.

10. C. subrobustum (Jorgensen) Nielsen

11. C. teres Kofoid

12. C. kofoidii Jorgensen

13. C. bohmii n. sp.

14. C. lineatum (Ehrenberg) Cleve

15. C. setaceum Jorgensen

ra. Subgenus AMPHICERATIUM

16. C. geniculat\mi (Lemmermann) Cleve

17. C. bigelowii Kofoid

18. C. inf latum (Kofoid) J6rgensen

19. C. longirostrum Gourret

20. C. falcatum (Kofoid) Jorgensen

21. C. extensum (Gourret) Cleve

22. C. fusus (Ehrenberg) Dujardin

IV. Subgenus EUCERATIUM

23. C. tripos (O. F. Muller) Nitzsch

subsp. atlanticum Ostenfeld
subsp. semipulchellum Jorgensen

24. C. pulchellum Schroder

25. C. humile Jorgensen

26. C. breve (Ostenfeld and Schmidt) Schroder

27. C. compressum Gran

28. C. euarcuatum Jorgensen

29. C. filicorne Nielsen

30. C symmetricum Pavillard

var. symmetricum (Pavillard)
var. coarctatum (Pavillard)
var. orthoceros Jorgensen

31. C. axiale Kofoid

32. C. aultii n. sp.

33. C. azoricum Cleve

34. C. petersii Nielsen

35. C. arietinum Cleve

subsp. arietinum (Cleve)
subsp. bucephalum (Cleve)
subsp. gracilentum (Jorgensen)

36. C. declinatum Karsten

37. C. gibberum Gourret

f. subaequale Jorgensen

38. C. concilians Jorgensen

var. subaequale n. var.

39. C. lunula Schimper

40. C. carnegiei n. sp.

41. C. contortum Cleve

42. C. limulus Gourret

43. C. paradoxides Cleve

44. C. platycorne Daday

45. C. ranipes Cleve

46. C. macroceros (Ehrenberg) Vanh5ffen

subsp. macroceros (Ehrenberg)
subsp. gallicum (Kofoid)

47. C. massiliense (Gourret) J5rgensen

48. C. deflexum (Kofoid) Jorgensen

49. C. carriense Gourret

50. C. contrarium (Gourret) Pavillard

51. C. trichoceros (Ehrenberg) Kofoid

52. C. vultur Cleve

var. vultur (Cleve)
var. japonicum (SchrSder)
var. sumatranum (Karsten)
var. pavillardii (Jorgensen)
var. regxilare n. var.
var. reversum n. var.
var. recurvum Jorgensen

53. C. horridum Gran

var. horridum (Gran)
var. moUe (Kofoid)
var. claviger (Kofoid)

54. C. tenue Ostenfeld and Schmidt

var. inclinatum (Kofoid) Jorgensen
var. tenuissimum (Kofoid) Jorgensen

55. C. longissimum (Schroder) Kofoid

56. C. arcticum (Ehrenberg) Cleve

var. arcticum (Ehrenberg) Cleve
var. longipes (Bailey) Gran
var. ventricosum Ostenfeld

57. C. hexacanthum Gourret

58. C. reflexum Cleve

Subgenus POROCERATIUM

1. Ceratlum praelongum (Lemmermann) Kofoid
Figure 1, chart 1, appendix table 1

A rare, strictly tropical species, confined to sta-
tions where the surface temperature was over 20° C.
Found in the Gulf Stream to where the surface tempera-
ture dropped to 20.°5 (station 15); also sporadically in
the Sargasso and Caribbean seas. Peters (1934) did not
find it farther south than 25° south latitude in the South
Atlantic. At Carnegie stations it was foimd as far north
as 38.°5 north (station 15).

In the Pacific Nielsen (1934) reported it at various
stations in the South Equatorial Current. Okamura

(1912) found it at Japan, but it was not reported from the
Pacific by Bohm (1931). In the southeastern Pacific, in
the Carnegie collections, it was found at one station in
the Himiboldt Current (station 69), at two stations south
of Easter Island, and at eight scattered stations in the
South Equatorial Current. It was found continuously in
the two series of stations across the equatorial currents
in the central and western Pacific; at one station between
Guam and Japan, at two stations off Japan, and at one sta-
tion off California. It was not found farther north than
latitude 35° north in the North Pacific, nor farther south
than 32° south in the South Pacific. It was notably ab-
sent from the loop of stations northeast of Hawaii and in
the Galapagos-Panama region.

Ceratiimi praelongum was never found in great num-
bers. In the Carnegie collection it was found only at forty
stations --seven in the Atlantic and thirty-three in the
Pacific. There were sixty-seven records of occurrence,
fifty-one of which were rare and sixteen occasional.
Forty-nine of the records were from net samples, eight-
een from pump samples.

Ceratlum praelongum Is a shade species according
to Nielsen (1934). In the Carnegie collection it was found
more frequently at 50 meters than at the surface or 100
meters, and more often at 100 meters than at the surface
(table 2). Thus, the Carnegie data tend to substantiate
Nielsen's classification of this species as a shade form.

CERATIUM SPECIES OF CARNEGIE COLLECTION

15

Table 2. Records of occurrence of C. praelongum
at three levels

Occur -

Depth in meter

s

renc6

50

100

A

B

A

B

A

B

Rare
Occasional

12
5

3.1 28 9.9
1.3 6 2.1

11 4.3
5 2.0

Total

17

4.4 34 12.0

16 6.3

A=Niunber of records. B=Per cent of total number of
samples collected at that depth.

Although C. praelongum was limited to warm water
it did not show any preference for oligotrophic water.
Only sixteen of the sixty -seven records were in water
containing less than 10 mg P04/m3. The surface tem-
peratures at the stations where it occurred, varied from
20.°4 to 29.°4 C. The ranges of environmental conditions
in situ were: temperature, 14.°2 to 29.°4 C; salinity, 33.4
to 37.1 per mille; pH, 7.87 to 8.47; phosphate, 2 to 63
mg P04/m3.

Remarks . - -Dissections were made of the plates of
C. praelongum . The pattern Is shown in ventral and left
lateral views in figures 1 A and B. It will be noted that
in this species two apical plates touch the ventral area,
whereas Jorgensen (1911) stated that in the subgenus
Poroceratium only the large flat ventral apical touches
the ventral area. Kofoid (1907b) devised anomenclature
for the plates of Ceratium. For Poroceratium he start-
ed nimibering the apical plates with the wide ventral
one, basing his system on the pattern of C. gravidum in
which this plate is the only one to touch the ventral area.
Thus he called this one the apical plate. Jorgensen (1911),
on the other hand, called it apical four. A knowledge of
the pattern of C. praelongum . a member of Porocera-
tium . shows that Jorgensen's system is preferable In
that it indicates homologies. The Suture in C. praelon-
gum . which separates the ventral and left apical plates,
is homologous with the ventral suture of C. tripos ,
which separates the first and fourth apicals, so that the
ventral plate of Poroceratium is homologous with the
fourth of C. tripos and should be designated apical 4
as in Jorgensen's system. Ceratium praelongum . in its
epithecal plate pattern, is thus transitional between
other members of Poroceratium and Euceratium.

Variation. - -There is considerable variation in the
shape of the body of C. praelongum but the variants
seemed to bear no relation to geographic location or en-
vironmental conditions. The forebody may be constrict-
ed just anterior to the girdle as in figure lA, or in the
middle region (fig. IC), or may be quite regular as in
figure ID. The last figure shows an individual with very
heavy hypothecal walls and thick antapical horns. None
of the variations found could be satisfactorily grouped
into varieties.

2. Ceratium cephalotum (Lemmermaxm) Jorgensen
Fig^ure 2, chart 2, appendix table 2

This is a rare, intolerant tropical species. Jorgen-
sen (1911) stated that it has been reported from the
North Atlantic only north of the West Indies; Peters
(1934) found it only at one station in the South Atlantic.
In the Carnegie collection it was found in the Atlantic at

two stations between the North Atlantic Drift and the
Sargasso Sea at 100 meters in both cases.

The species is never found in great numbers, fii
the Carnegie collection it was found at a total of thirty-
two stations; two in the Atlantic, thirty in the Pacific.
There were thirty-nine records of occurrence- -thirty-
six rare and three occasional. Thirty-six of the records
were from the net samples; three from the pump sam-
ples.

Ceratium rephalntiim is a shade species according
to Nielsen (1934), and this classification is substantiated
by the Carnegie data as shown in table 3. It was found
in an increasing percentage of the samples with increase
in depth. Although Nielsen found it only in the samples
taken between 100 to 50 meters, in the Carnegie collec-
tion it was found almost as frequently at the surface as
at 50 meters.

Table 3.

Records of occurrence of C. cephalotum
at three levels

Occur-

Depth in meters

rence

50

100

A

B

A 1 B

A B

Rare
Occasional

12

1

3.2 9 3.2
0.2 2 0.7

15 5.9
....

Total

13

3.4 11 3.9

15 5.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

The Carnegie data suggest that this species prefers
oligotrophic water. It is always found in warm water.
The surface temperatures of the water at the stations
where it was found varied from 23.°6 to 29.°4C. Twenty-
four of the thirty-nine records were in water containing
less than 10 mg P04/m3. It is particularly noteworthy
-in this connection that the species was found m a contin-
uous series of stations north and east of Hawaii and all
are included in the great "desert" of the North Pacific.
Peters' (1934) record for the South Atlantic was also in
oligotrophic water. The ranges of environmental condi-
tions m_situ were: temperature, 13.°8 to 28.°7C; salinity,
34.1 to 36.5 per mllle; pH, 8.17 to 8.47; phosphate, 4 to
36 mg P04/m3.

Ceratium rep halntinn is not very variable. Some-
times the overgrowth of the right side is less pronounced
(fig. 2B).

3. Ceratium gravidimi Gourret
Figures 3 and 4, chart 3, appendix table 3

This is a relatively rare, but widespread, slightly
tolerant tropical species found at most of the Carnegie
tropical stations. Peters (1934) found It only in warm
oligotrophic water in the South Atlantic. In the Carnegie
collection it was found at most (seventeen) of the twenty-
three stations in the warm Atlantic region. It was not
found at any Atlantic station where the surface tempera-
ture was less than 20° C. All the Atlantic stations were
in oligotrophic water with phosphate content less than 10
mg at the surface, except at station 1 where it was thirty-
two. The temperature there, however, was 24°.

In the Pacific Nielsen (1934) found C. gravidum in
both eutrophic and oligotrophic water at scattered stations

16

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

in the South Pacific. In the Carnegie collection it was
pretty well scattered throughout the tropical waters of
the Pacific, occurring at seventy-five stations and
sometimes in water of somewhat lower temperature
than that of the Atlantic. Off Japan it occurred to 40.°5
north, surface temperature 15.° 9 C (station 117). Off
Calif ornia it occurred to 34 ° north, surface tempera-
ture 19.°1 C (station 31). In the southeastern Pacific it
occurred south of Easter Island to 34 ° south, surface
temperature 19° C (station 57). In the extreme south-
eastern Pacific it was not common, occurring at only
four stations east of the line of stations in the longitude
of Easter Island.

Although widespread, this species is never very
abundant. There were 182 records of occurrence, 144
of which were rare and 38 occasional. Of these records,
164 were from the net samples and 18 from the pump
samples.

Ceratium gravidum was classified as a shade spe-
cies by Nielsen (1934). In oligotrophic water he found
that it had its main distribution in the levels from 200 to
100 meters, although he found it in the upper levels in
eutrophic water. The Carnegie data substantiate this
classification of Nielsen. The total number of records
(in percentage) at 50 meters is more than three times
that at the surface; the number of 100-meter records
shows an increase over that for 50 meters (see table 4).
This distribution is shown also by the rare and occasion-
al records separately. Thus, C. gravidum is a decided
shade species.

Table 4.

Records of occurrence of C. gravidimi
at three levels

Occur-

Depth in meters

rence

50

100

A

1 B

A 1 B

A 1 B

Rare
Occasional

27
3

7.7 58 20.5 59 23.0
0.8 14 3.6 21 8.3

Total

30

8.5 72 24.1 80 31.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

In the Carnegie collection as a whole, C. gravidum
seemed to be found in both oligotrophic and eutrophic
water. Sixty-one of the 182 records of occurrence were
in water containing less than 10 mg P04/m3. The sur-
face temperatures at the stations where it occurred,
varied from 15.°9 to 29.°5C. The ranges of environmen-
tal conditions in situ were: temperature, 10.°6 to 29.°3 C;
salinity, 33.4 to 37.0 per mille; pH, 7.76 to 8.44; phos-
phate, 2 to 153 mg P04/m3.

Variation. -- Ceratium gravidmn is a variable spe-
cies. Jorgensen (1911) divided it into three varieties and
later (1920) described a fourth variety. These have been
more or less accepted by later workers (Nielsen, 1934
and Peters, 1934), although apparently no one hitherto
has had sufficient material to test the validity of these
varieties. In the Carnegie studies hundreds of specimens
were examined, and it was evident that no taxonomlc
grouping could be made of the wide variations in body
form found in this species. Representative specimens
are shown in figures 3 and 4. Each of these drawings is
representative of a group of individuals examined and
other variations between these were found. Thus, there

is a complete intergradation between the wide rotund
form shown in figure 3A (which is even wider than JOr-
gensen's var. latum) and the narrow form shown in fig-
ure 4U (which, in turn, is narrower than JOrgensen's
var. angustum ). The characteristics of the antapical
horns such as length, thickness, curvature, and diver-
gence (which have also been used in differentiating the
varieties) also vary in such a manner that no varietal
segregations are indicated. An attempt was made to
correlate the variations of C. gravidum with geographic
location, or with different water masses. No correlation
was found; the different variations were indiscriminately
scattered throughout the areas investigated. Peters
(1934) found the different varieties in the same regions
of the South Atlantic. Nielsen (1934) found var. latum to
be restricted to the eutrophic water of the eastern part
of the South Equatorial Current, whereas var. elegans
and angustum were found there as well as in the oligo-
trophic western part and in the East Australian Current.
Nielsen's results are not corroborated by the Carnegie
collections. M the latter, specimens as wide or wider
than var. latum were, indeed, found in the eutrophic
water in the region of the Galapagos Islands, but they
also occurred in the oligotrophic water of the central
part of the North Equatorial Current.

Subgenus BICERATIUM

4. Ceratium digitatimi Schutt
Figure 5, chart 4, appendix table 4

This is a rare intolerant tropical species, one of
the most rare of the Carnegie collection. B is apparent-
ly restricted to warm water as it never approached the
20° surface isotherm except in the region of Easter Is-
land. It is also remarkable that it was never found near
large land masses, although Jorgensen (1920) foimd it in
the Mediterranean. Peters (1934) found it at only four
stations in the South Atlantic in his "main distribution
region" of warm oligotrophic water. On the Carnegie
cruise it was found only at two stations in the Atlantic,
in the North Equatorial Current.

Nielsen (1934) is the only previous author to report
C. digitatum from the Pacific. He foxind it at three wide-
ly separated stations in the South Equatorial Current. In
the Carnegie collection it was found at twenty-three
widely separated stations, from Easter Island to Guam
and to a point midway between Hawaii and California.
The most northern station is 33.°5 north; the most south-
ern, 31.° 5 south. On the German South Polar expedition
C. digitatum was found to 31 ° south. This seemed extra-
ordinary to Peters (1934) who did not find it farther south
than 20° south on the Meteor expedition. He explained the
fact by stating that it was probably carried south in the
Agulhas Current. The Carnegie southern records in the
southeastern Pacific may thus be explained by a drift
from the northeast, in the large vortex characteristic of
that region. The absence of this species off Japan, how-
ever, where it should be carried by the extension of the
North Equatorial and Japan currents, and in the North At-
lantic Drift, are noteworthy and can only be explained on
the assumption that the species is so rare that its detec-
tion anywhere in the ocean must be considered fortuitous.
Its limits of distribution, thus, will not be well defined
until much more survey work has been done.

Ceratium digitatum is always found singly or in small

CERATIUM SPECIES OF CARNEGIE COLLECTION

17

numbers. In the Carnegie collection there were twenty-
six records of its occurrence--all rare. Eighteen of
these records were from net samples, eight from pump
samples.

This species is undoubtedly a shade species, al-
though previously insufficient data have been at hand to
demonstrate this. JSrgensen (1920) foimd it in the Med-
iterranean at the deeper levels in the summer but attri-
buted this occurrence to another cause (see p. 12).
Nielsen (1934) considered it a shade species, although
'lis few data were from above the 50-meter level. Since
two of his three records for this species were from sta-
tions rich in plankton, however, the occurrence above
this level does not necessarily indicate that it is not a
shade species. The Carnegie records substantiate Niel-
sen's classification of this species as a shade species.
It was found more frequently with increase in depth
(table 5).

Table 5.

Records of occurrence of C. digitatum
at three levels

Occur -

Depth in meters

r6nc6

J

50

100

A

B

A 1 B

A B

Rare

6

1.5 8 2.1 12 4.7

Total

6

1.5 8 2.1 12 4.7

A=Number of records. B=Per cent of total number of
samples collected at that depth.

It is doubtful if C. digitatum prefers oligotrophic
water. Ten of the twenty-six Carnegie records were
from water containing less than 10 mg P04/m^. This
is not owing to the fact that most of the records are
subsurface records. The phosphate content of the sur-
face water at the stations where the species was foimd
shows the same distribution. Ten of these are below
10 mg and the rest above, with some as high as 42. The
surface temperatures at the stations where C . digitatum
occurred, varied from 22.°4 to 29.°4 C. The ranges of
environmental conditions in situ were as follows: tem-
perature, 12.°5 to 28.°5 C; salinity, 34.3 to 36.5 per
mille; pH, 8.16 to 3.39; phosphate, 3 to 46 mg P04/m3.

Variation. --Very few variations have been reported
for this species, probably owing to scanty material.
Jorgensen (1920), however, described a variant, var.
rotundatum . found in the Mediterranean and also in the
Guinea Current. It is characterized by more rounded
apex, less bent apical horn, and somewhat larger size.
Peters (1934) stated that in the South Atlantic he found
specimens with more rounded epitheca than Jorgensen's
variety, fo the Carnegie collection two specimens were
found which answer well to Jorgensen's var. rotxmdatum .
They occurred in the surface samples frbm stations 52
and 54, both south of Easter Island in temperatures of
22.°5 and 23.°4C (figs. 5A and B). a is probable that this
form represents a species distinct from C. digitatum as
no intergrades were found between it and the typical
form. The typical C. digitatum does not vary greatly,
although the length of the left antapical horn varies
somewhat in different specimens (figs. 4C and E).

5. Ceratium candelabrum (Ehrenberg) Stein
Figure 6, chart 5, appendix table 5

This is a common, slightly tolerant tropical spec:es.
It is not confined to warm water. At Carnegie stations
the limits of distribution paralleled rather closely the
surface isotherm of 15° C in all three regions where the
lines of stations extended into colder regions, namely,
in the southeastern Pacific, off Japan, and off the west
coast of the Americas. These limits of distribution are
very significant, inasmuch as within these limits there
are extremely few stations without records of occur-
rence. At six stations off Peru it was not found, although
the surface temperatures there were above 19° C. Ap-
parently the peculiar environmental conditions in this
region have an inhibitory effect on C. candelabrum . In
the Atlantic the species occurred almost continuously in
the warmer water, but its northern limit there coincided
with the surface isotherm of 20° C rather than of 15° as
in the Pacific. In respect to latitude, C. candelabrum
was found approximately to 40° south and 40° north in
the Pacific, and to 42° north in the Atlantic. Its south-
ern limit in the South Atlantic is also at about 40° south
(Peters, 1934). Some of the stations of the Dana expedi-
tion extend, in the southwestern Pacific, beyond 40° south.
The southern records of C. candelabrum , however, end
at the fortieth parallel. This roimds out a remarkable
correlation between latitude and the distribution of a
Ceratium. It must be mentioned, however, that in the
southwestern Pacific, as well as in other parts of the
Pacific, the latitudes of 40° are closely paralleled bythe
surface isotherms of 15° C.

This species is one of the most common of the genus.
In the Carnegie collection it was found at 136 stations --
26 in the Atlantic and 110 in the Pacific. There were 353
records of occurrence--206 rare, 128 occasional, and 19
common. Of these, 243 were from net samples and 110
from pimap samples.

In regard to vertical distribution, Neilsen (1934)
classified this species as a surface form. This classifi-
cation is substantiated by the Carnegie observations. As
can be seen in table 6, C. candelabrum was found most
frequently at 50 meters (44.2 per cent of the 50-meter
samples). It was found almost as often (36.0 per cent of
the surface samples) at the surface, but only half as fre-
quently at 100 meters (21.0 per cent). It is more signifi-
cant that the records of occasional and common occurred
much more frequently at the surface and 50 meters than
at 100 meters.

Table 6.

Records of occurrence of C. candelabrum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

75

57

8

18.4 81

13.4 42

2.1 5

28.4
14.2

1.7

50

29

6

20.0
0.8
0.2

Total

140

33.9 128

44.3

85

21.0

=Number of records. B=Per cent of total number of
samples collected at that depth.

18

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Cpratiiim ranHplahniTn was found throughout the

warm-water regions without respect to the nutrient con-
tent of the water. Thus, 117 of the 353 records of oc-
currence were in water containing less than 10 mg
P04/m3. The range of surface temperatures at the sta-
tions where it occurred was from 15.°9 to 29.°5 C. The
ranges of environmental conditions in situ were: tem-
perature, 8f8 to 29.°5 C; salinity, 30.0 to 37.1 per mille;
pH, 7.76 to 8.47; phosphate, 3 to 189 mg P04/m3.

Variation. - -Several varieties of C. candelabrum
have been described, the most important of which are
var. depressum Jorgensen (1920) and f. commune Bohm
(1931). Nielsen (1934) distinguished these two Variations
and stated that both are found in the western part of the
Pacific investigated by him but only var. depressum was
found in the eastern Pacific. The writers are inclined
to agree with Peters (1934) that the variations in this
species are too continuous for any separation of varie-
ties, although it must be stated that most of the Carnegie
material resembled the var. depressum form. Figure 6
shows a series of specimens with variously bent horns,
both apical and antapical, and with various lengths of
horns.

6. Ceratium furca (Ehrenberg) Dujardin
Figure 7, chart 6, appendix table 6

Ceratium furca Is an interesting cosmopolitan spe-
cies probably composed of many subspecies or races.
Jorgensen (1911) designated two subspecies; a northern
form, subsp. a berghii and a tropical form, 6 eugram-
minn. Ceratium hircus Schroder, which Jffrgensen ac-
cepted as an independent species, is probably also a va-
riety of C. furca. In his study of Mediterranean Ceratla,
Jorgensen (1920) designated many more varieties. Bohm
(1931) attempted to show that in his material from the
western Pacific there were two genotypes, both belonging
to the tropical subspecies eugrammum . These were sep-
arated on the basis of total length; a short variety with
length from 130 to 168 microns, and a large varietyfrom
170 to 244 microns. Nielsen (1934) did not consider
Bohm's evidence conclusive, and made a statistical study
of length in his material from the South Pacific. In only
one sample did he find a separation into two groups on
this basis. Bi other samples long ones or short ones
were represented, and often medium-sized ones. Never-
theless Nielsen considered that two races were repre-
sented.

As a result of the study of the Carnage material the
writers tend toward JOrgensen's (1920) opinion, that the
species is a complex of many varieties and races, not of
two. No separation could be made on the basis of size
alone. No attempt was made to separate the different
variants. The number is great and the intergradations
common, fii figure 6 several variants are shown.

The distribution of C. furca is of particular interest.
Peters (1934), in his study of the Ceratia of the Meteor
expedition in the South Atlantic, concluded that C. furca
is a typically neritic species. It was found abundantly in
the mouths of great rivers and in regions of upwelling. It
seemed to avoid the middle regions of the oceans. Niel-
son (1934) accepted this classification of the species and
stated that it is distinctly a neritic form, although also
found at many oceanic stations. He found it at every sta-
tion but two between Panama and Tahiti, and at various
stations between that point and Australia. He stated that

its occurrence at these oceanic stations can be explained
by the disturbed water about the islands in the western
Pacific and by the eutrophic water occurrence in the
eastern part of the South Equatorial Current. Such a
broad use of the term "neritic," however, leaves no
room for the term oceanic so such usage cannot be ac-
cepted. Nielsen's data show, on the other hand, that C^
furca is oceanic as well as neritic. In the Carnegie col-
lection the species did show a tendency to occur more
frequently and more abundantly in richer water but,
nevertheless, was often found in oligotrophic water.
Only 40 of the 231 records of occurrence were in water
containing less than 10 mg P04/m^.

In the Carnegie collection there is no indication of
an avoidance of oceanic water or of the middle regions
of the open oceans on the part of C. furca . Not only are
there many station records for the open North Atlantic
and the Pacific, but there are as many records of occa-
sional and common for stations remote from land as for
stations close to land (see chart 6).

In the Atlantic there were two distributional areas,
probably representing the distributions of the two sub-
species; a northern one from stations 6 to 11a (from
southwest of Ireland around the North Sea and Iceland to
southeast of Greenland), and a tropical region from sta-
tions 14 to 24 (straight down through the middle of the
Atlantic). There was also one station in the Caribbean.
In the Pacific, also, the distribution was interrupted.
The species was found quite regularly in the southeast-
ern Pacific, but was scattered irregularly over the other
tropical regions of the Pacific. It was notably absent
from the northern line of stations from stations 118 to
128 inclusive. No northern form was found in this ocean.

The surface temperatures in this " C. furca -free"
water in the northern Pacific varied from 7.° 2 to 16.°4C.
This should not have constituted a barrier to this species
as it was found In the Atlantic where the surface temper-
atures were as low as 8.°9. The tropical subspecies was
confined to warm water in the Atlantic (surface temper-
atures 21.°2 to 28.°0 C), but in the Pacific it extended into
somewhat cooler water (surface temperatures from 15.°0
to 29.°4 C). Obviously the northern subspecies of the
Atlantic was not found in the Pacific.

Ceratium furca is frequently found in quantities.
Records of "abundant" were from the North Sea, south-
eastern Pacific, and southeast of Japan. The species was
found at 91 Carnegie stations--19 in the Atlantic and 72
in the Pacific. There were 232 records of occurrence,
with 140 rare, 65 occasional, 22 common, and 4 abundant.

Nielsen (1934) classified C. furca as a surface form.
The Carnegie data show that it is truly not a shade form,

Table 7.

Records of occurrence of C. furca
at three levels

Depth

in meters

r6nc6

50

100

A

B

A

B

A

B

Rare

Occasional
Common
Abundant

68

36

14

4

17.7
9.5
3.6
1.4

42

14

3

14.9
4.9
1.6

30

15

5

11.8
5.9
1.8

Total

122

32.2

59

21.4

50

ly.5

A=Number of records. B=Per cent of total number of
samples coUected at that depth.

CERATIUM SPECIES OF CARNEGIE COLLECTION

19

but is found at depths below the surface in a great many
cases. The table of frequencies (table 7), however,
shows that it was found most frequently at the surface.
This is particularly true of the records of "common"
and "abundant."

The ranges of environmental conditions in situ were
as follows: temperature, 6.°6 to 29.°4 C; salinity, 33.1 to
37.0 per mille; pH, 7.63 to 3.47; phosphate, 3 to 233 mg
P04/m3.

7. Ceratium belone Cleve
Figure 8, chart 7, appendix table 7

This is a rare, intolerant tropical species confined
to water of high temperature. Peters (1934) found the
species in the South Atlantic not beyond 30° south. Niel-
sen (1934) found it in the Pacific scattered over the en-
tire region of the South Equatorial Current.

In the Carnegie collection C. belone was foxmd at
twenty-six stations; two in the Atlantic and twenty-four
in the Pacific. The Atlantic records were in the middle
North Atlantic where the surface temperatures were
about 24° C (station lb) and 27.°0 C (station 18). The
most northern of these was at 38° north latitude.

In the Pacific C. belone was found at one station off
Peru, at eleven stations around and north of Easter Is-
land, at seven stations east and north of Samoa, at one
station in the North Equatorial Current, and at three
stations in the central North Pacific (chart 7). Its lati-
tudinal limits were 33f 5 north and 32.°0 south, which is
a rather limited distribution for a species of Ceratium.
It is a curious and unexplainable fact that the species
was not found west of 179° west longitude. The surface
temperatures at the Pacific record stations varied from
20.° 4 to 29.°4 C. K must, therefore, be considered a
stenothermal species. Although it always occurred in
water of high temperature, it was seldom found in water
ef extremely low phosphate content. Only six of the fifty-
one records of occurrence were from water containing
less than 10 mg P04/m3.

The ranges of environmental conditions in situ were:
temperature, 16.°0 to 29.°4 C; salinity, 34.1 to 37.0 per
mllle; pH, 8.11 to 8.47; phosphate, 5 to 48 mg P04/m3.

Nielsen's (1934) classification of C. belone asasur-
face species is substantiated by the Carnegie data (table
8), although there were two records of "occasional" at
100 meters. The total percentages, however, show a
much more frequent occurrence at the surface than at the
lower levels.

The occurrence of C. belone is sporadic. Unlike
some of the rarer tropical species, when it does occur,
it is sometimes foimd in considerable numbers. Peters
(1934) found this to be true in the South Atlantic. Like-
wise in the Carnegie collection there were several rec-
ords of "occasional." The total number of records was
fifty-one, of which forty-four were rare and seven were
occasional.

Variation. --Ceratium belone is rather constant in
shape and size, except for the divergence of the antapi-
cal horns and the length of the apical horn. There is an
extreme variation in the total length of the specimens
(from 335 to 910 microns), but this difference is owing
entirely to the difference in the length of the apical horn.
The length of the hypotheca is constant. In only one
specimen (fig. 8H) was this elongate. The diameter, d,
varied from 20 to 30 microns.

Table 8.

Records of occurrence of C. belone Cleve
at three levels

Depth in meters

rence

50

100

A

1

B

A

B

A B

Rare
Occasional

23
4

6.0 10 3.5
1.0 1 0.4

11 4.3
2 0.8

Total

27

7.0 11 3.9

13 5.1

A=Nimiber of records. B=Per cent of total number of
samples collected at that depth.

8. Ceratiimi incisum (Karsten) JSrgensen
Figure 9, chart 8, appendix table 8

This is a rare, strictly tropical species with limited
distribution. Peters (1934) found it at only four stations
in the South Atlantic, and Nielsen (1934) found it at scat-
tered stations in the South Equatorial Current of the Pa-
cific. In the Carnegie collection it occurred at twenty-
seven stations- -two in the Atlantic and twenty-five in the
Pacific. There were thirty records of occurrence, all
rare. Twenty-two of these were from net samples and
eight were from pump samples.

Of the two Atlantic stations, one was in the open At-
lantic at latitude 36.° 5 north, longitude 46.° 5 west (station
16); the other was in the Caribbean Sea. M the Pacific
the stations were scattered: one off Ecuador, six north
of Easter Island at about latitude 10° south, twelve be-
tween Samoa and Guam, four in the central crossing of
the equatorial regions, and two north of Hawaii. It was
very restricted in its latitudinal distribution and was ab-
sent from the southeastern Pacific region except at one
station off Ecuador. Its latitudinal limits in the Pacific
were 29° north and 15° south. In the Atlantic it occurred
to 36.°5 north.

Ceratium incisum is a stenothermal form similar to
C. belone. The surface temperatures at stations at which
it occurred in the Atlantic were 25.°9 and 28.°5 C. In the
Pacific the range was 24.°6 to 29.°5 C. The ranges of en-
vironmental conditions in situ were: temperature, 14.°2
to 29.°3 C; salinity, 33.7 to 36.3 per mille; pH, 7.87 to
8.39; phosphate, 4 to 48 mg P04/m3.

Apparently C. incisum is at home in oligotrophic
water. Ten of the thirty records of occurrence were in
water containing less than 10 mg P04/m3. Nielsen
(1934) classified this species as a surface species. In
the Carnegie collection it was found about equally at the
three levels (table 9).

Table 9. Records of occurrence of C. incisum
at three levels

Depth in meters

pence

50

100

A

B

A { B

A 1 B

Rare

11

2.9 9 3.2 10 3.9

Total

11

2.9 9 3.2 10 3.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

20

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

9. Ceratium pentagonum Gourret
Figures IOC, D, H-N, chart 9, appendix table 9

This is a cosmopolitan species composed of three
subspecies: a widespread tropical form, subsp. tenerum
(Jorgensen); a subantarctic form, subsp. robustimi
(Cleve); and a northern Pacific form, subsp. pacificum
n. subsp. The authors agree with Nielsen (1934) that
Jorgensen' s var. subrobustum is specifically distinct
(see below).

Subspecies tenerum is a widespread tropical form
somewhat more common than C. subrobustum . but with
a similar distribution except that it seems to be some-
what more tolerant of colder water. It was found at
eighty-one stations- -fifteen in the Atlantic and sixty-
eight in the Pacific. There were 189 records of occur-
rence: 152 rare, 34 occasional, and 1 common. Of
these records, 73 were from net samples, 114 from
pump samples. Expressed in percentages these are:
13.6 per cent of net samples, and 31.8 per cent of pump
samples. It is thus evident that the tropical form of C^
■ jentagonum is too small to be properly collected in the
tow nets.

The subspecies was found in the Atlantic mostly in
the central line of stations; it was absent from the
warmer part of the Gulf Stream and Caribbean Sea. In
the Pacific its distribution is somewhat irregular, but
the records are probably incomplete there, owing to the
fact that the distinctness of C. subrobustum was not re-
alized at the time of the routine census of the samples,
so some of its records probably belong to C. pentagonum.
Ceratium pentagonum . however, extended beyond the
range of C. subrobustum in the southeastern Pacific and
off California.

The surface temperatures at the stations at which
subsp. tenerum occurred, varied from 15.°0 to 29.°3 C.
The ranges of environmental conditions in situ were:
temperature, 13.°1 to 29.''3 C; salinity, 33.4 to 37.0 per
mille; pH, 7.76 to 8.47; phosphate, 3 to 233 mgP04/m3.

Nielsen (1934) listed C. pentagonimi as a surface
species. In the Carnegie collection it was found less
frequently at 100 meters than at the surface and 50-
meter depths, and slightly more frequently at 50 meters
than at the surface. Thus it cannot be classed as a
shade species. Ceratium subrobustum . on the other
hand, is definitely a shade species, with its greatest fre-
quency at 100 meters (cf. tables 10 and 11).

Table 10. Records of occurrence of C. pentagonum
subsp. tenerum at three levels

Depth in meter

s

50

100

A

1 B

A

B

A

B

Rare

Occasional

Common

56
18

1

14.5
4.6
0.2

56

11

19.1
4.0

40 15.3
5 1.6

Total

75

19.3

67

23.1

45 16.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

Ceratium pentagonum is a remarkable species, in
that it has never been collected from the cold northern
Atlantic although it is represented in the subantarctic
by a cold-water subspecies. In the Carnegie collection
the distribution of the species in the North Atlantic

terminated under 45° north latitude (station 13a), which
is somewhat farther north than previous authors had
found it. The surface temperature there, however, was
above 20° C. It apparently cannot endure transfer into
temperatures much below this value.

The conspicuous absence of C. pentagonum in the
northern Atlantic was particularly noted by Peters (1934)
who mapped the distribution of subsp. robust\im in the
subantarctic waters of the South Atlantic. Peters em-
phasized the unequal distribution in which the species is
represented in the southern cold water by the most south-
erly member of the genus, but is entirely absent in the
cold waters of the North Atlantic. Subspecies robustum
doubtless populates the entire subantarctic region as
there are no barriers in this region. Thus, it probably
occurs in the southern latitudes of the Pacific. Peters
did notfind it in the Atlantic north of about 48°, however,
except where it was displaced along the South American
coast by the Falkland Current to about 32° south. This
subspecies was not found in the Carnegie collection prob-
ably because the stations were not far enough south.
The most southern station (station 60) was at about 40°
south. It is probable that the subspecies is displaced
northward in the Humboldt Current to 30° or so, but in
these latitudes the Carnegie stations were too far from
the coast to detect it.

In view of this knowledge it was of particular inter-
est to search for this species in the cold waters of the
North Pacific. The investigations here revealed the
presence of a cold-water subspecies peculiar to this re-
gion, subsp. pacificum n. subsp.

A series of forms was found south of the Aleutian
Islands and they have been grouped together imder this
subspecies. The extreme form (fig. 101) is remarkably
different from the other subspecies of C. pentagonum .
It is large (transdiameter 78 microns), with heavy walls
covered with thick ridges and lists, and the antapical
horns are long (equal to the transdiameter or longer).
All the forms in this cold-water region, however, do not
conform to this description, and in the west grade into
the southern form (see figs. IOC and J). Another varia-
tion is shown in figure lOD. These intergrades indicate
that the extreme form is of subspecific rank only.

With the finding of this subspecies it was evident that
in the case of C. pentagonum there is a bipolarity in the
Pacific with subsp. robustum in the antarctic and subsp.
pacificum in the north, whereas in the Atlantic only the
southern form is represented. This distribution consti-
tutes the most convincing evidence of the planktological
isolation of the North Pacific waters (see p. 11).

Subspecies pacificum was found at seven stations
and in seventeen samples (chart 9). The surface tem-
peratures of the stations where it occurred, varied from
7.°2 to 10.°5 C. The ranges of environmental conditions
in situ were: temperature, 4.°2 to 10.°5 C; salinity, 32.7
to 32.9 per mille; pH, 7.98 to 8.03; phosphate, 103 to 175
mg P04/m3.

10. Ceratiimi subrobustum (JSrgensen) Nielsen
Figures lOA, B, E-G, chart 10, appendix table 10

This is a rare, intolerant tropical species. Peters
(1934) found this form in the South Atlantic only in the
Falkland Current and for that reason considered it a
subspecies, raising it from the varietal rank given by
Jorgensen (1920). Nielsen (1934) found it in the Pacific

CERATIUM SPECIES OF CARNEGIE COLLECTION

21

in the eastern part of the South Equatorial Current and
at Australia, but not at the colder stations. He raised it
to the rank of an independent species, and separated it
from C. pentagonum on the basis of size of body and
thickness of walls. He gave the transdiameter of C.
subrobustum as 77 to 89 microns, and that of C. penta -
g onum up to 72 microns. In the Carnegie material the
transdiameter of C. robustum varied from 72 to 90 mi-
crons; those of C . pentagonum (excluding subsp. pacificum
n. subsp.) were from 38 to 66 microns. The transdiam-
eter of subsp. pacificum was 55 to 78 microns.

Ceratium subrobustimi is probably rarer than C.
pentagonum but, nevertheless, it is much more wide-
spread than previously was supposed. In the Carnegie
collection it occurred at forty-five stations--two in the
North Atlantic, where it had not been found before, and
forty-three in the Pacific. There were eighty-one rec-
ords of occurrence: sixty-three rare, fifteen occasion-
al, and three common. Of these records, fifty were from
net samples, and thirty-one were from pump samples.
Unfortunately, when the routine examination of this spe-
cies was conducted, the independence of this species was
not realized, so the records of its occurrence are not
complete. For this reason, the peculiar clumping of the
stations in the Pacific must be considered artificial and
not representing exactly the actual distribution.

It may be noted that the species is a warm -water
form, however, not extending into water of low tempera-
ture. The surface temperatures at the stations where it
occurred, were all above 20° C except in the southeastern
Pacific, where it was found at stations with surf ace tem-
peratures as low as 17° C. The ranges of environmen-
tal conditions in situ were: temperature, 14. °3 to 28.° 5
C; salinity, 34.0 to 39.8 per mille; pH, 7.17 to 8.39;
phosphate, 3 to 64 mg P04/m3.

Nielsen (1934) stated that in the Dana collections C.
subrobustum was found only in the upper 50 meters.
The Carnegie data are at variance with this and strongly
suggest that the species is a shade species. As can be
seen from table 11, the species was foundmore frequent-
ly with increase in depth to 100 meters. Particularly
significant are the records of "occasional," which were
found in 0.3 per cent of the surface samples, 2.1 per
cent of the 50-meter samples, and 3.2 per cent of the
100-meter samples. In this respect C. subrobustum dif-
fers from C. pentagonum . which was found more fre-
quently at 50 meters.

Table 11.

Records of occurrence of C. subrobustimi
at three levels

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional
Common

12

1
1

2.8
0.3
0.3

16
6

1

5.7
2.1
0.4

35
8
1

13.0
3.2
0.4

Total

14

3.4

23

8.2

44

16.6

A=Number of records. B=Per cent of total number of
samples collected at that depth.

11. Ceratium teres Kofoid
Figures IIB-D, chart 11, appendix table 11

This is a rare, slightly tolerant tropical species,
although widespread. to the Carnegie collection it

occurred at sixty-two stations — twelve in the Atlantic,
and fifty in the Pacific. There were 106 records of oc-
currence, with 85 rare and 21 occasional. The species
is so small that it was collected more fre4uently in the
pump samples. There were 30 net records and 76 pump
records. Expressed in percentages these are 5.6 per
cent of the net samples, 20.8 per cent of the pump sam-
ples.

In regard to geographic distribution, the species is
rather too irregular to draw any conclusions. Its ab-
sence is particularly noticeable, however, in the region
of Panama, Galapagos, and southward. It occurs, on the
other hand, at Easter Island and south to 40° south.

Ceratium teres was mostly confined to warm water.
In the Atlantic the surface temperatures at its record
stations were all above 24.°8 C; in the Pacific, above
20.°6 C, except in the southeastern region where it was
as low as 15.°0 C (station 60). The ranges of environ-
mental conditions in situ were: temperature, ll.°4 to
29.°4 C; salinity, 33.9 to 37.1 per mille; pH, 7.99 to 8.39;
phosphate, 3 to 123 mg P04/m3.

Ceratium teres does not avoid oligotrophlc water.
Forty-seven of the 106 records of occurrence were in
water containing less than 10 mg P04/m3.

Nielsen (1934) found this species only in the sam-
ples taken from 50 to meters. In the Carnegie collec-
tion it was found about as frequently in the 100-meter
samples as in the samples from higher levels (table 12).
The surface contamination of the deeper nets, however,
must be taken into consideration here.

Table 12.

Records of occurrence of C. teres
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A 1 B

A B

Rare
Occasional

36

10

9.1 31 11.0 18 7.1
2.6 3 1.1 8 3.2

Total

46

11.7 34 12.1 26 10.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

12. Ceratium kofoidll Jorgensen
Figure IIH, chart 12, appendix table 12

This is a very rare, slightly tolerant tropical spe-
cies, seldom collected because of its extremely small
size. In the Carnegie collection it was found at eight
stations in the Pacific. There were twelve records of
occurrence, ten of which were rare and two occasional.
It was found predominantly in the pump samples because
of the smaller mesh of the pump fUter net. Two of the
records were from net samples and ten were from pump
samples.

The record stations for C. kofoidii were widely
scattered (chart 12). The surface temperatures at the
stations where It occurred, varied from 16.°2 to 27f 7 C.
The environmental conditions in situ were: temperature,
ll.°7 to 27.°6 C; salinity, 34.4 to 35.3 per mille; pH, 8.03
to 8.34; phosphate, 5 to 58 mg P04/m3.

Six of the records were from surface samples, three
from 50-meter samples, and three from the lOO-meter
samples. Thus, it is probably a surface species. Four
of the twelve records were from water containing less
than 10 mg P04/m3.

22

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

13. Ceratium bohmii n. sp.
Figure 12, appendix table 13

Dimensions: diameter, d, 20.5 (18-24) microns.

This species is closely related to C. kofoidii with
which Bohm (1931, figs.9C, E, and F) confused it. The
dimensions are about the same. The body, however, is
much more elongated and the horns, particularly the
antapicals, are much longer. The length of the hypothe-
ca from the base of the right antapical horn to the pos-
terior edge of the girdle is more than 0.5 d, whereas in
C. kofoidii this length is less than 0.5 d. The right ant-
apical diverges laterally only slightly or not at all.

Ceratium bohmii is a tropical Pacific species, prob-
ably slightly tolerant, although this cannot be determined
at present because of the paucity of records for the spe-
cies. It was found at six stations widely scattered as
follows: two between Ecuador and the Galapagos, two
north of the Phoenix Islands, and two off Japan. There
were twelve records of occurrence — eight rare and four
occasional. The species was collected more frequently
with the pump, probably because of its small size.
There were three net records and nine pump records.

The surface temperatures at the stations where the
species was found, varied from 18.°7 to 27.°9 C. The en-
vironmental conditions in situ were: temperature, 13.°9
to 27.°9 C; salinity, 33.7 to 35.3 per mille; pH, 7.85 to
8.22; phosphate, 4 to 161 mg P04/m3.

Ceratium bohmii is probably a surface species, al-
though there are not sufficient records to establish this.
Five of the records were from surface samples, three
were from 50-meter samples, and four from 100-meter
samples.

14. Ceratium lineatimi (Ehrenberg) Cleve
Figures llE-G, chart 12, appendix table 14

Ceratium lineatum is a rare species which is diffi-
cult to classify geographically. It occurs in the cold
North Atlantic region and at three stations in the warm
Atlantic (chart 12). In the Pacific it occurred only off
Japan (stations 115a to 117). It is possibly a subpolar
species. Peters (1934) found it in the South Atlantic,
mostly in the cooler waters. At Carnegie Atlantic sta-
tions the species was found mostly in colder waters with
surface temperatures from 10.°9 to 15.°5 C except at sta-
tions 17 and 19. At the Pacific record stations the sur-
face temperatures varied from 15.°9 to 16.°1 C. The
environmental conditions in situ were: temperature, 6.° 7
to 21.°2 C; salinity: 33.8 to 37.0 per miUe; pH, 7.98 to
8.34; phosphate, 3 to 99 mg P04/m3.

The total number of station records was ten, of which
seven were in the Atlantic and three in the Pacific. There
were twenty-six records of occurrence, ten of which were
rare, fourteen occasional, and two common. The records
of common were from off Japan. Twenty-one of the rec-
ords were from net samples and five were from pump
samples. The species was found frequently at the sur-
face, with thirteen records for the surface, six records
for 50 meters, and seven recrods for 100 meters.

The authors were unable to separate C . minutum Jor-
gensen from C. lineatum. Jorgensen (1920) distinguished
C. minutum from C. lineatum by its smaller dimensions,
less robust theca, comparatively shorter and broader body,
shorter antapical horns, and relatively shorter right ant-
apical horn. Carnegie specimens with the shape of C. minu-

tum have relatively thick walls (fig. IIF); specimens with
the typical shape of C. lineatum are in the size range of C.
minutum . i.e., transdiameter 25 to 28 microns (figs. HE, G).

15. Ceratium setaceum JOrgensen
Figure HA, appendix table 15

This is a very rare, slightly tolerant tropical spe-
cies. Nielsen (1934), who first reported it from the Pa-
cific, found it at only two stations- -one north of Samoa,
and the other west of New Zealand. In the Carnegie col-
lection it was found at only two stations also --station 70,
off Peru, and station 110, between Guam and Japan. Judg-
ing from the scattered nature of these records, it is
probable that C. setaceum is a widespread tropical spe-
cies, but occurs in such sparse numbers that it is seldom
collected. Another reason for its reported scarcity is
undoubtedly its small size which enables it to pass
through many nets. This is demonstrated by the fact that
all the Carnegie records were from the pump samples
which were collected by a much finer (no. 20) cloth than
the tow nets. The records were at and 50 meters at
station 70, and 100 meters at station 110.

The surface temperatures at the above two stations
were 21. °2 and 23.°9 C. The ranges of environmental con-
ditions in situ were: temperature, 15.°4 to 21. °2 C; salin-
ity, 34.7 to 35.1 per mille; pH, 7.88 to 8.14; phosphate,
11 to 178 mg P04/m3.

Subgenus AMPHICERATIUM

16. Ceratium geniculatum (Lemmermann) Cleve
Figure 11 J

This is a very rare tropical species. Only one spec-
imen was found in the Carnegie collection, and this was
partially broken (fig. IIJ). It occurred in the 50 -meter
pump sample at station 45, latitude 4.°5 south, longitude
105° west. The temperature was 22.°4 C; salinity, 35.2
per mille; pH, 8.13; and phosphate 46 mg P04/m*.

17. Ceratium bigelowii Kofoid
Figures 11 1, K-M, chart 13, appendix table 16

This is a very rare intolerant tropical species, ap-
parently confined to the Pacific and Indian oceans. It has
never been reported from the Atlantic. It was reported
from the Pacific by Kofoid (1907a) and Nielsen (1934),
and from the Pacific and Vidian oceans by BOhm (1931).
Nielsen foimd it at four stations in the South Equatorial
Current. The Carnegie records contribute considerably
to our knowledge of the distribution of this species in the
Pacific. They show that it is not restricted to any par-
ticular current system, but is distributed over the warm
tropical waters in both hemispheres. &i the Carnegie
collection it was found at twenty-four stations (chart 13).
These are distributed in the southeastern Pacific to lati-
tude 34° south (station 57), in the northwestern Pacific to
latitude 23.°5 north, and in the northeastern Pacific to
26° north latitude. There are forty records of occur-
rence --thirty- six rare and four occasional. The records
of occasional were at 50 meters at stations 95, 96, and
159, and also at the surface at the last station. All three

CERATIUM SPECIES OF CARNEGIE COLLECTION

23

of these stations were in the region north of Samoa, and
it is possible that this is a region usually rich in this
species since one of these stations, station 159, was oc-
cupied seven months after the other two.

All the records of C.bigelowii are from warm water
and are remote from land. The surface temperatures at
the stations where it was found, varied from 20.° 7 to
29.°4 C, except at station 57 south of Easter Island where
the surface temperature was 19. °0. The environmental
conditions in situ were: temperature, 14.°3 to 29°3 C;
salinity, 34.2 to 36.4 per mille; pH, 8.10 to 8.39; phos-
phate, 3 to 50 mg P04/m3.

According to Nielsen's theory (1934), C. bigelowii
should be a shade species since it has an expanded flat-
tened body. Since there have been so few records of this
species, however, it has not been possible to examine
this feature heretofore. The Carnegie records in this
connection are not conclusive. The records of rare and
the total number of records for the three levels show an
increase In frequency with increase in depth (table 13).
The records of occasional, however, stand in opposition
to this, with the greatest frequency at 50 meters. Wheth-
er C. bigelowii is a shade species cannot be decided def-
initely imtil more records of its occurrence have been
accumulated.

Since it is a large species, it was collected more
often in the net, with thirty-four net records, and only
three pump records.

Table 13.

Records of occurrence of C. bigelowii
at three levels

Occur-

Depth in meters

50

100

A

B

A

B

A

B

Rare
Occasional

9
1

2.1

0.3

13 3.9
3 1.0

14 5.5

Total

10

2.4

16 4.9

14 5.5

A=Number of records. B=Per cent of total number of
samples collected at fhat depth.

18. Ceratium inf latum (Kofoid) JOrgensen
Figures llO-S, chart 14, appendix table 17

This species is very closely related to the two fol-
lowing. Jorgensen (1920) considered C. inf latum (Kof.)
Jorgensen, C. longiro strum Gourret, C. falcatum Kofoid,
and C. falcatiforme Jorgensen all separate species,
whereas Peters (1934) treated them as members of a
Formenkreis with subspecific rank. Nielsen (1934) con-
sidered them distinct. Whatever the taxonomic value of
these groups, there is certainly an intergradation be-
tween them. In this report the first three have been kept
separate, but this arrangement must be considered ten-
tative until more statistical work is done on the group.
It was found impossible, however, to separate C.falcati -
forme from C. falcatum. Therefore, only the first three
species are listed in this report.

One of the characters which Jorgensen (1920) used
for the differentiation of the above species was length of
epitheca relative to length of hypotheca--the ep/hyp ratio.
For each species he gave the mean, minimum, and maxi-
mum ep/hyp ratios. It should be noted that his fractions for
maxima represent the maximum length of epitheca over
the maximum length of hypotheca and not the maximum
ep/hyp ratios that follow them, which might be inferred

from his method of presentation. The same holds true
for his minima. Unfortunately, many of Jorgensen's fig-
ures for this group indicate ratios which are beyond the
range that he states in the text. For instance, his fig-
ures of C. longiro strum (figs. 26 and 27) show ep/hyp
ratios of 1.525 and 1.55, whereas in the description he
states that the ma;>Lmum ratio is 1.45. Which is correct,
the figures or the description? Owing to these inaccura-
cies it is difficult to evaluate Jorgensen's data.

A study of the Carnegie specimens, however, re-
sulted in measurements showing fairly close agreement
with those of Jorgensen. For C. inf latum he gave an
ep/hyp ratio of 1.15 (1.06 to 1.26) for an a form and 1.21
(1.18 to 1.23) for "another form." This ratio in the
present material was 1.11 (0.86 to 1.25). Equally close
agreement was found in the case of the other species
(which see). The total length in our material is 1010
microns (640 to 1460); the diameter 40 microns (30 to
80). Incidentally, C. inf latum in our material is easily
separated from C. longirostrum bv its diameter alone,
since the latter has a diameter of 19 microns (15 to 27).

It must also be noted that C. bigelowii is closely re-
lated to C. inf latum and should be included in the group
in case it is treated as a Formenkreis. In the Carnegie
material quite wide specimens of C. inf latum were found
which were difficult to separate from C. bigelowii . This
was particularly true where there was a lateral flatten-
ing of the body (fig. IIR).

Ceratium inflatum is a rare, intolerant tropical spe-
cies confined entirely to warm water. In the Carnegie
collection it occurred at twenty-four stations- -five in the
Atlantic and nineteen in the Pacific. There were thirty-
one records of occurrence, thirty of which were rare and
one occasional. Twenty-five were from net samples and
six were from pump samples.

The record stations for this species were scattered
over the warm-water regions. All of them were remote
from land, except one near Panama. The surf ace tempera-
tures at these stations were from 26.°0 to 27.°2 in the At-
lantic, and from 22.°3 to 28.°0C in the Pacific, except at
one station south of Easter Island where the temperature
was 19° C. The environmental conditions in situ were:
temperature, 16.°6 to 28.°6 C; salinity, 34.3 to 36.8 per
mille; pH, 8.00 to 8.37; phosphate, 3 to 121 mg P04/m3.

Ceratium inflatum is apparently an oligotrophic spe-
cies. Not only was it foiud principally in warm water re-
mote from land, but It was usually found in water of low
nutrient content. More than half (seventeen) of the
twenty-nine records of its occurrence were in water con-
taining less than 10 mg P04/m3.

Nielsen (1934) suggested that this was a surface spe-
cies although also found in the deeper samples. In the
Carnegie collection it had its maximum frequency at 50
meters (see table 14). Thus, the data acquired so far in-
dicate that the occurrence of C. inflatum is rather gener-
al in a vertical direction.

Table 14.

Records of occurrence of C. inflatum
at three levels

Occur-

Depth in meters

rence

50

100

A B

A B

A

B

Rare
Occasional

Total

12

2.9

11
1

3.9
0.4

2.4

12

2.9

12 4.3

2.4

A=Number of records. B=Per cent of total number of
samples collected at that depth.

24

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

19. Ceratium longiro strum Gourret
Figures IIT-V, chart 14, appendix table 18

Total lengtii, 644 (570-730) microns; diameter, 19
(15-27) microns; ep/hyp ratio, 1.36 (1.19-1.50). Jorgen-
sen (1920) gave the ep/hyp ratio as 1.38 (1.26-1.45).
Thus, there is close agreement between the world mate-
rial of the Carnegie and the Mediterranean material of
Jorgensen.

Ceratium longirostrum is a rare, intolerant tropi-
cal species with a distribution similar to that of C. in-
f latum (see chart 14). In the Carnegie collection it was
found at sixteen stations — two in the Atlantic and four-
teen in the Pacific. There were twenty-six records of
occurrence — twenty-one rare and five occasional.
Eighteen records were from net samples and eight were
from pump samples.

The surface temperatures at the stations where it
occurred, varied from 22.°3 to 29.°3 C, except at one
station (station 2) in the North Atlantic Drift where the
temperature was 20f 5 C. The environmental conditions
in situ were: temperature, 21.°6 to 28.°5 C; salinity, 34.4
to 36.6 per mille; pH, 8.14 to 8.37; phosphate, 5 to 36mg
P04/m3.

Not only is C. longirostrum found in warm water
remote from land, but it is often in water poor in nutri-
ents. Fourteen of the twenty-six records of occurrence
were in water containing less than 10 mg P04/m3.

Nielsen (1934) listed C. longirostrum as a surface
species. In the Carnepie collection there were insuffi-
cient records to draw any conclusions in this regard. It
was found least often at 50 meters, but more frequently
at 100 meters than at the surface (table 15).

Table 15. Records of occurrence of C. longirostrum
at three levels

Depth in meters

r6nc6

50

100

A

B

A

B

A

B

Rare
Occasional

10
2

2.6
0.5

3 1.1 8 3.1
2 0.7 1 0.4

Total

12

3.1

5 1.8 9 3.5

A=Number of records B=Per cent of total number of
samples collected at that depth.

20. Ceratium falcatum (Kofoid) Jorgensen
Figures IIW-AA, chart 15, appendix table 19

Total length, 513 (350-750) microns; diameter, 23
(15-35) microns; ep/hyp ratio, 1.38 (1.00-0.72). Jorgen-
sen (1920) gave the ep/hyp ratios for C. falcatum as 1.56
(1.38-1.70) and for C. falcatiforme . 1.13 to 1.20. The
Carnegie range thus includes both of these. It was im-
possible to separate in the Carnegie material a small
form answering to C. falcatiforme Jorgensen. Jorgensen
stated that the length of this form was about 280 microns
which is smaller than any Carnegie specimen measured.
Many specimens were found, however, which had the
shape of the species in question, although larger than
this. These inter graded into the typical C. falcatum (figs.
IIW-AA). It is probable that C. falcatiforme is simply
an extreme of the species C. falcatum. Nielsen (1934)
considered it a well-defined species, but presented no

measurements, ratios, or other evidence which might
indicate that it is distinct in the Pacific. He simply
stated that it is "small."

Ceratium falcatum is a rare, intolerant tropical
species with a distribution similar to that of C. inflatxmi
and C. longirostrum (see chart 15). In the Carnegie
collection it was found at thirty stations- -one in the At-
lantic and twenty-nine in the Pacific. There were forty-
seven records of occurrence--forty-slx rape and one
occasional. Thirty-four records were from net sam-
ples and thirteen were from pump samples.

The surface temperatures at the stations where it
occurred, varied from 19.°0 to 28.°4 C. The environ-
mental conditions in situ were: temperature, 14.°2 to
28.°5 C; salinity, 34.5 to 36.4 per mille; pH, 7.86 to 8.39;
phosphate, 4 to 198 mg P04/m3.

Although this species was found mostly in oiigotroph-
ic water, this was not so evident as in the case of C. in-
flatum and C. longirostrum. Only ten of the forty-seven
records were in water containing less than 10 mg
P04/m3.

Nielsen (1934) listed C. falr.ahim and C. falcatiforme
as surface species. This is borne out by the Carnegie
data (see table 16).

Table 16.

Records of occurrence of C. falcatum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare
Occasional

Total

26
1

5.7
0.3

10

2.8

10

3.9

27 6.0

10 2.8

10 3.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

21. Ceratiimi extensum (Gourret) Cleve
Figures IIBB-DD, chart 16, appendix table 20

This species includes C. strictum (Okamura and
Nishikawa) Kofoid, which is characterized by the pres-
ence of a well-formed right antapical horn. Since the
presence of this horn is not a reliable character, and
since it is the only one separating C. strictum from C.
extensum . it is no longer considered a separate species
(see Jorgensen, 1920; Peters, 1934; and Nielsen, 1934).

The species is a widespread very tolerant tropical
form. In the Carnegie collection it occurred at 104 sta-
tions--14 in the Atlantic and 90 in the Pacific. These
were distributed in the various regions as follows: 3 in
the cold North AUantic (in the North Atlantic Drift), 11
in the warm Atlantic, 1 in the cold North Pacific (off
California), 79 in the warm Pacific, and 10 in the south-
east Pacific. There were 234 records of occurrence —
136 rare, 81 occasional, and 17 common. Of the total
number, 144 were from net samples and 90 were from
pump samples.

The surface temperatures at the stations where it
occurred, varied from 12.°4 to 29.°4 C. In the northeast-
ern Pacific and in the southeastern Pacific it extended
into water with surface temperatures of 16.°4 and 14.°97
respectively. In the North Atlantic Drift it extended close

CERATIUM SPECIES OF CARNEGIE COLLECTION

25

to the British Isles to a surface temperature of 12.°4 C.
The environmental conditions in situ varied as follows:
temperature, 10.°4 to 29.°4 C; salinity, 29.7 to 37.1 per
mille; pH, 8.0 to 8.39; phosphate, 2 to 99 mg P04/m3.
Nielsen (1934) classified C. extensum as a surface
species. The Carnegie data corroborate this classifica-
tion. The species was found with decreasing frequency
with Increase in depth (see table 17). If due correction
were made for contamination of the open nets, this fea-
ture would be more pronounced.

Table 17.

Records of occurrence of C. extensum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

72
43

9

18.7

11.2

2.3

33

28
6

11.7
9.9
2.1

30

10

2

11.8
3.9
0.8

Total

124

32.2

67

23.7

42

16.5

A=Number of records. B=Per cent of total number of
samples collected at that depth.

22. Ceratium fusus (Ehrenberg) Dujardin
Figures llEE, 13A-D, chart 17, appendix table 21

Ceratium fusus is a widespread cosmopolitan spe-
cies. It may be composed of subspecies but so far no
morphological differences warranting a division of the
species have been found. Jorgensen (1920) believed that
he could distinguish a tropical subspecies (seta [Ehren-
berg]) from the northern form of the species, but later
workers have not been able to verify this (Peters, 1934;
Nielsen, 1934). Peters found several different forms in
different parts of the South Atlantic, but theyintergraded
so much that he was not able to separate them satisfac-
torily. Nielsen considered the species to represent a
single genotype which is definitely modified as to size by
ecological conditions. His data for the South Pacific
show an increase in size under neritic conditions.

In the Carnegie collection a statistical difference
was found between the forms growing in the different life
zones (see table 18). It is evident from these data thatt
the specimens from cold-water regions have a distinctly
greater diameter than those from the warm-water re-
gions as shown by both the means and extremes. The
length, however, does not show the same correlation.
The specimens from the warm Atlantic showed the great-
est average length, whereas those from the warm Pacific
had the smallest. It is to be hoped that a finer morpho-
logical analysis of this species will throw some light on
the taxonomic-ecologic problems involved.

In the Carnegie collection C . fusus was found at ninety-
four stations — twenty-five in the Atlantic, and sixty-nine
in the Pacific. These stations were distributed from the
warmest to the coldest regions visited (see chart 17).
There are particular Unes of stations, however, where
C. fusus was not found. These distributional gaps cannot
be explained on the basis of general rarity of the species.
In the Carnegie collection there were 194 records of its
occurrence.. Of these, 109 were rare, 67 occasional, 11
common, and 7 abundant. Most of the records of higher
frequency, however, were from the cold Atlantic region.

Table 18. Size variation in Ceratium fusus

Region

Total length
in microns

Diameter
in microns

Cold

Atlantic
Warm

410 (310-500)

21.6 (20-28)

Atlantic
Cold North

459 (345-434)

20 (16-25)

Pacific
Warm

420 (330-520)

24 (20-32)

Pacific
Southeast

329 (168-422)

15 (10-25)

Pacific

345 (250-405)

17 (12-22)

and it should be noted that the lines of stations where
the species was not foimd are mostly in regions of low
phosphate content. Only 23 of the 194 records of occur-
rence were in water containing less than 10 mg P04/m3.
Thus, it is possible that C. fusus does not normally de-
velop to any great extent in oligotrophic water.

Since the species is cosmopolitan, it was foimd in a
great range of conditions. The surface temperatures at
the stations where it occurred, varied from 7.°2 to 29.°5
C. The environmental conditions in situ were: temper-
ature, 2.°1 to 29.°5 C; salinity, 32.7 to 37.0 per mille; pH,
7.71 to 8.37; phosphate, 4 to 209 mg P04/m3.

The very narrow body of C. fusus permits it to es-
cape from the larger meshed nets. Consequently there
are relatively more pump records for the species than
net records. It was found in only 18 per cent of the net
samples, but in 26 per cent of the pump samples.

Nielsen (1934) classified C. fusus as a surface spe-
cies. This classification is probably correct. In the
Carnegie collection the number of records decreased
with increase in depth (see table 19). After correcting
for upper -level contamination in the open nets, it is ob-
vious that the species is much more frequent in the sur-
face layer.

Table 19.

Records of occurrence of C. fusus
at three levels

Occur-

Depth in meters

rence

50

100

A

B

A

B

A

B

Rare

44

10.9

34

12.0

31

12.2

Occasional

37

9.4

21

7.5

9

3.5

Common

8

2.8

1

0.4

2

0.7

Abundant

4

1.4

2

0.8

1

0.4

Total

93

24.5

58

21.7

43

16.8

A=Number of records. B=Per cent of total number of
samples collected at that depth.

Subgenus EUCERATIUM

23. Ceratium tripos (O. F. Muller) Nitzsch

No species of the genus is so complex and variable
as C. tripos . For a time there was a tendency to sepa-
rate the components of this form into separate units with
specific rank. This movement was led by JOrgensen

26

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

(1911, 1920). Later a tendency developed to lump these
units into a Formenkrelsor a few species. Peters (1934)
included under C. tripos all the varieties of pulchellum
(see Jorgensen, 1920). He divided the species into three
subspecies: atlanticum . pulchellum. and semipulchellum .
Nielsen (1934), on the other hand, recognized four sepa-
rate species in this group as follows: C. tripodoides . C.
semipulchellmn . C. pulchelltun . and C. tripos . He sepa-
rated the first three species from each other on the ba-
sis of statistical studies of individual plankton samples.
He found a grouping according to diameter and "antapi-
cal length." The last, C. tripos , which he found only in
the cold water in the region of Australia and New Zea-
land, he considered distinct because of its shorter apical
horn and stronger antapicals. These characters can
hardly be sufficient for the sepai'ation of a species in
this variable group. It is probable that the New Zealand-
Australian form is simply a southern subspecies or race.
Certainly Nielsen's drawings (figs. 32 and 33) do not
show sufficient distinctness for a separation from his C.
semipulchellum (fig. 30).

The present authors lean toward the view of Peters
(1934) in regard to this group, with the exception that C^
pulchellum (eupulchellimi of Jorgensen) has been treated
as a separate species. Considering the collection as a
whole, there was very little inter gradation between this
form and semipulchellum . Between semipulchellum and
tripos , on the contrary, there are many intergradations.
i.e., in various characters such as size, spread of horns,
and antapical curvature. It was considered advisable for
the present to treat these two units as subspecies. They,
in turn, are each composed of an endless number of va-
rieties or races, which, however, are not recorded here
as they are Impossible to separate with our present
knowledge of morphological characters in this genus.

Ceratiimi tripos subsp. atlanticum (Ostenfeld)
Figures 13E-K, chart 18, appendix table 22

This subspecies is more characteristic of colder
waters although it is sometimes found in the tropics;
The tropical forms are probably separate races; some
of them vary toward the semipulchellum form.

In the Carnegie collection subsp. atlanticum was
found throughout the cold Atlantic region and at ten sta-
tions in the warm Atlantic region. In the Pacific it was
found in the cold North Pacific region to the west and
east but not in the most northern part. In the warm Pa-
cific region it was found at ten stations northof 27° north
latitude, and at two stations in the southeastern part. In
the southeast Pacific region it was found at thirteen sta-
tions. The total number of record stations for the sub-
species was sixty-one — twenty-six of which were in the
Atlantic and thirty-five in the Pacific.

It was not found so often as subsp. semipulchellum
but a high percentage of the records were above "rare."
The total number of occurrences was 163, of which 49
were rare, 61 occasional, 45 common, and 8 abundant.

Since the subspecies is cosmopolitan, the range of
environmental conditions was great. It was not, however,
foimd at the wannest stations of the Pacific. The range
of surface temperatures at the record stations in the At-
lantic was from 8.°9 to 28.°5C; in the Pacific from 6.°9 to
27.°1 C. The environmental conditions In situ were: tem-
perature, 6.°1 to 28.°2 C; salinity, 31.7 to 36.4 per miUe;
pH, 7.80 to 8.37; phosphate, 2 to 181 mg P04/m3.

This subspecies, like the rest of the Formenkreis,
is a surface species, although it is also found in consid-
erable numbers In the deeper levels. As shown in table
20, subsp. atlanticum was found most frequently in the
surface samples.

Table 20. Records of occurrence of C. tripos
subsp. atlantictun at three levels

Occur-

Depth in meters

rence

50

100

A

B

A

B

A

B

Rare

22

5.7

14

4.6

13

5.1

Occasional

32

8.3

13

4.6

16

6.3

Common

25

6.5

11

3.9

9

3.5

Abundant

7

1.8

1

0.4

....

Total

86

22.3

39

13.5

38

14.9

A=Number of records. B=Per cent of total niimber of
samples collected at that depth.

Ceratium tripos subsp. semipulchellum (Jorgensen)
Figures 13L-N, chart 19, appendix table 23

This subspecies is a slightly tolerant tropical form
which occurs almost continuously in the warm regions
and the southeast Pacific region (chart 19). It is limited
to the tropical waters except in the southeastern Pacific
and northeast of Japan. In the Atlantic it was limited en-
tirely to the warm Atlantic region. In the Pacific it oc-
curred at one station northeast of Japan in the cold North
Pacific region, although not in water of low temperature;
the lowest surface temperature being 16.°1 C. In the
southeastern Pacific region it was found at all but three
stations. The surface temperatures at the record sta-
tions there were as low as 15° C. The total range of
surface temperatures was from 15° to 29.°5 C. The
ranges of environmental conditions In situ were: tem-
perature, ll.°4 to 29f4 C; salinity, 31.6 to 36.6 permUle;
pH, 7.76 to 8.47; phosphate, 2 to 198 mg P04/m3.

The subspecies was found at a total of 127 stations--
21 in the Atlantic and 106 in the Pacific. It is one of the
most common of the Ceratia. This is shown, not only by
the large number of stations at which it was found, but
also by the large number of single records of occurrence
and relative abundance. There were 391 records of oc-
currence, with 161 rare, 156 occasional, 72 common, and
2 abundant. Since it is a large form, it was found more
often in the net samples. There were 256 net records
and 135 pimip records.

The subspecies definitely does not avoid oligotrophlc
water, as its record stations are unbroken through re-
gions of poor water, and 140 of the 391 records of occur-
rence were In water containing less than 10 mg P04/m3.

In regard to depth, Melsen (1934) states that seml-
pulchellum is a definite surface form. The Carnegie
data, on the contrary, indicate that it lives throughout the
upper 100-meter layer. The greatest percentage of
"rare" records was at 100 meters (see table 21); of
"occasional" records at the surface; of "common" rec-
ords at 50 meters; and the greatest total percentage was
at 50 meters. The large number of records for the 50-
and 100-meter levels cannot be explained entirely on the
basis of contamination of the open nets since there were
twenty-three stations scattered over the Atlantic and

CERATIUM SPECIES OF CARNEGIE COLLECTION

27

Pacific where the subspecies was not recorded for the
surface but was recorded for either or both the 50- and
100-meter levels.

Table 21. Records of occurrence of C. tripos
subsp. semipulchellum at three levels.

Occur-
rence

Depth

in meters

50

100

A

B

A

B

A

B

Rare

Occasional
Common
Abundant

60
72
28

1

15.6

18.7

7.2

0.2

46

47

31

1

16.0

16.6

11.0

0.3

55

37

13

21.6

14.6

5.1

Total

161

41.7

125

43.9

105

41.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

24. Ceratium pulchellum Schroder

(=C. pulchellum f. eupulchellum Jorgensen)

Figures 14B-F, chart 20, appendix table 24

As stated above (p. 26), C. pulchellum Schroder is
considered distinct from the C. tripos-C. semipulcheU-
lum complex. It is characterized by short antapical
horns and small size. The diameter in our material was
46 (22-60) microns.

Ceratium pulchellum is a fairly common, slightly
tolerant tropical species. In the Carnegie collection it
was found at sixty-nine stations- -eleven in the Atlantic
and fifty-eight in the Pacific. In the Atlantic the stations
were restricted to the warm Atlantic region; in the Pa-
cific they were scattered over the warm Pacific region
and the southern part of the southeast Pacific region.
The total number of sample records for the species was
158, of which 97 were rare, 47 occasional, and 14 com-
mon. It is a curious fact that the relative number of
pump records for the species is greater than the net
samples. The species was found in 15 per cent (82) of
the net samples and 21 per cent (76) of the pump sam-
ples. This probably is owing to the relatively small di-
ameter of the species and to the short antapical horns,
which permit the species to pass more readily through
the coarser cloth of the tow net.

The surface temperatures at the stations where the
species was found, varied from 16.°9 (in the southeastern
Pacific) to 29.°5 C (north of Samoa). The environmental
conditions in situ were: temperature, ll.°4to 29.°4 C;

Table 22. Records of occurrence of C. pulchellum
at three levels

Occur -

Depth in meters

50

100

A

B

A

B

A B

Rare

Occasional

Common

47

32

9

12.2
8.3
2.3

27 9.5
9 3.2
3 1.1

23 9.1
6 2.4
2 0.8

Total

88

22.8

39 13.8

31 12.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

salinity, 34.0 to 37.1 per miUe; pH, 7.76 to 8.39; phos-
phate, 3 to 123 mg P04/m3.

Ceratium pulchellum is decidedly a surface species
as can be seen from table 22.

25. Ceratium humile Jorgensen
Figure 14A

This species was reported from Japanese waters by
JOrgensen (1911), and from the Australian region by Niel-
sen (1934). The latter author considered it endemic in
the East-Asiatic Australian region. He also considered
it a purely neritic species. For this reason, the single
record of this species in the Carneg ie collection is re-
markable. 1 was foimd in the surface sample at station
1 in the Gulf Stream . This not only extends the known
distribution of the species to another ocean but indicates
that, at least in the Atlantic, it is not necessarily a neri-
tic species. The surface temperature at this station was
24.°0 C. The specimen (fig. 14A) resembles those figured
by Jorgensen (1911, figs. 82 and 83) somewhat more than
that figured by Nielsen (1934, fig. 34) as the right horn is
more ascending.

26. Ceratium breve (Ostenfeld and Schmidt) Schroder
Figures 14G-P, chart 21, appendix table 25

This is avariable species which exhibits transitions
to many other species of the genus, namely, C. schmidtii .
C. tripos , and C. arcuatum . Peters (1934) has sugge_sted
that it be included in the Formenkreis C. tripos . Bohm
stressed the similarity between C. breve and C. arcuatum
Cleve and the present authors believe that C. schmidtii
likewise is similar or identical to C. breve. A series of
specimens is shown in figures 14G to P which extends
from the form with short, curved antapicals as in C.
schmidtii (fig. 141) to the open parallelum type simulat-
ing C. arcuatum (fig. 14K). It is probable, as Bohm (1931)
stated, that the transition to C. arcuatum is superficial
rather than genetic. Whether C. schmidtii should be in-
cluded in C. breve is more difficult to say. According to
Jorgensen (1911) the body of C. schmidtii is concave so
the antapical horns have a ventral curvature. This fea-
ture, however, is sometimes found in C. breve , so it is
not possible to separate these two forms on that basis.

Ceratium breve is an intolerant tropical species. In
the Carnegie collection it was found at ninety-two stations- -
eighteen in the Atlantic and seventy-four in the Pacific.
All the Atlantic stations were in the warm Atlantic region.
The Pacific stations were in the warm Pacific region or
the western part of the southeast Pacific region, with the
single exception of station 70 off Peru. The surface tem-
perature there, however, was high, namely, 21.°2C. The
species was not found in most of the southeastern Pacific
nor in most of the area between California and Hawaii.
This distribution may represent the effects of subpolar
currents (see p. 9). The complete range of surface tem-
peratures was 18.°7 to 29.°5 C. The environmental condi-
tions in situ were: temperature, 12^5 to 29.°4 C; salinity,
29.7 to 36.8 per mille; pH, 7.68 to 8.47; phosphate, 2 to
189 mg P04/m3.

Melsen (1934) classified C. breve as a surface form.
The Carnegie data corroborate this classification. As
shown in table 23, it was found less frequently in the
deeper samples. If due correction were made for the

28

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

contamination of the open nets this tendency would be
more pronounced.

Table 23. Records of occurrence of C. breve
at three levels

Occur-

Depth

in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

43
55
23

10.4

13.3

6.0

37

30
10

13.9

10.6

3.5

32

23
5

12.0
8.0
2.0

Total

121

29.7

77

28.0

60

22.0

A=Number of records. B=Per cent of total number of
samples collected at that depth.

The species is often found in comparatively large
numbers. There were 258 records of occurrence, of
which 112 were rare, 108 occasional, and 38 common.
Since it is a large species, it was found more often in the
net samples. There were 181 net samples and 77 pump
samples. Seventy -five of the records were in water con-
taining less than 10 mg P04/m3.

27. Ceratium compressimi Gran
Figure 15A

This is a very rare species previously known only
from the temperate waters of the North Atlantic. In the
Carnegie collection a single specimen was found. This
occurred in the 100-meter net sample from station 61 in
the southeastern Pacific, latitude 38.°5 south. Probably
this is a bipolar species occurring in the cooler waters
of both hemispheres. The surface temperature at station
61 was 16.°9 C. The environmental conditions at 100 me-
ters were: temperature, 10.°8 C; salinity, 34.0permille;
pH, 8.03; phosphate, 80 mg P04/m3.

28. Ceratiimi euarcuatum Jorgensen
Figures 15N-M, chart 22, appendix table 26

This is a distinct and easily recognized species
which varies comparatively little. It is an intolerant
tropical species, although found almost continuously in
the warm regions of the oceans (chart 22). It was found
at eighty-five stations- -fourteen in the Atlantic, and
seventy-one in the Pacific. In the Atlantic it was con-
fined to the warm Atlantic region and in the Pacific to
the warm Pacific region except for two stations in the
western part of the southeast Pacific region. The sur-
face temperatures at the stations where it occurred in
the Atlantic varied from 24.°0 to 28.°0 C and in the Pacif-
ic from 19.°0 to 29.°5 C. It was absent from the entire
southeastern Pacific east of 100° west longitude except
for one station west of the Galapagos Islands. Nielsen
(1934), however, foimd a single specimen in the Panama
region.

Ceratiimi euarcuatum probably is one of the best in-
dicators of warm tropical water. It apparently is of no
value as an indicator of intrusions of water of tropical
origin, however, since it does not seem to extend beyond
the tropical boundaries in such currents as the Kuroshio
and Gulf Stream. Apparently it does not endure a lowering

of the temperature. On the other hand, it is found within
the tropics in regions where cooler water has been
brought in from other regions or from vertical disturb-
ances. Thus, it was found west of the Galapagos where
the surface temperature was 19.°6 and south of Easter
Island where it was 19.°0 C. Nielsen (1934) found that it
was restricted to water where the surface temperatures
were above 22.° 5 C. During the cruise of the Dana the
temperatures in the region of the Galapagos were some-
what higher than when the Carnegie visited there but the
species was not found in this area by Nielsen. It is
probable that this region is in the line of distributional
limit for the species, that normally it does not occur
southeast of there, and that its occurrence in the region
of the Galapagos depends on the amount of mixing of the
waters in that region and on the amount of extrusion of
the water of the Humboldt Current. Peters (1934) con-
sidered C. euarcuatum an indicator of warm oligotrophlc
water in the South Atlantic. Nielsen (1934), on the other
hand, found it in water rich in nutrients and plankton as
well as in oligotrophlc water.

One might conclude from the Carnegie distribution
of C. euarcuatum in the Atlantic and North Pacific that
the species preferred oligotrophlc water because its dis-
tributional limits in these oceans parallel quite closely
the line of 10 mg P04/m3 (cf . charts 22 and 50). This
correlation, however, does not hold in the South Pacific
where the species was frequently found in water rich in
nutrients. Thus, we must conclude that it is not high
concentration of nutrients which is limiting the distribu-
tion of the species in the northern waters but some other
factor associated with it. Only 86 of the 185 records of
occurrence were in water containing less than 10 mg
P04/m3.

Within the tropics the species is fairly common so
there were many records of occurrence, namely, 185; of
which 97 were rare, 75 occasional, and 13 common. It
was found more often in net samples, with 128 net rec-
ords ana 57 pump records.

The environmental conditions in situ were: temper-
ature, 13.°6 to 29.°3 C; salinity, 33.9 to 37.1 per mille;
pH, 7.17 to 8.47; phosphate, 3 to 123 mg P04/m3.

Nielsen (1934) classified C. euarcuatum as a shade
species and the Carnegie data substantiate this classifi-
cation. As shown in table 24, it was fovind more frequent-
ly at the lower levels than at the surface.

Table 24.

Records of occurrence of C.
at three levels

euarcuatum

Occur-
rence

Depth in meters

50

100

A

B

A

B

A B

Rare

Occasional

Common

22

11

2

5.7
2.8
0.5

49

43

5

17.4

15.3

1.7

26

21

6

12.0
8.0
2.0

Total

35

9.0

97

34.4

53

22.0

A=Number of records. B=Per cent of total number of
samples collected at that depth.

29. Ceratium fllicorne Nielsen
Figures 15B-C, chart 23, appendix table 27

This species was described by Nielsen from the
Dana collection in the South Pacific. He stated that it is
closely related to both C. euarcuatum and C. svmmetricum

CERATIUM SPECIES OF CARNEGIE COLLECTION

29

and probably has been confused by some with the latter
species, although the original figures of Pavillard (1905)
and Jorgensen (1911, 1920) were not C. filicorne. In the
Carnegie collection this species was identified without
any difficulty, and no intergrades were foimd between it
and either C. euarcuatxun or C. svmmetricum .

Ceratium filicorne has a distribution of particular
interest. In the Carnegie collection it occurred only in
the South Pacific. This distribution is unique for the
Ceratium species. Since the only other records for the
species are those of Nielsen (which are also from the
South Pacific), its known distribution is limited to that
region. Nielsen's records run from the Tuamotu Islands
to Fiji. In the Carnegie collection there are three sta-
tions for the species between the Tuamotus and Samoa,
i.e., in the same general region of those of Nielsen; the
other stations, fifteen in number, are in the southeast-
ern Pacific. These, however, are limited to the warm
Pacific region, although they Include all the stations (4)
In the eastern extension of the region.

This is one of the most extraordinary distributions
found in the genus. If it were an extremely rare species
this distribution would not be remarkable as it might be
explained on the basis of incomplete sampling. This
probably is not the explanation in this case, however,
since the species was not found In particularly small
numbers within its range. Of the twenty-seven records
of occurrence only slxwere rare, whereas fourteen were
occasional, and seven were common. Fifteen of the rec-
ords were from net samples and twelve were from pump
samples. Why the species is not found in the equatorial
regions is an enigma. Is there something peculiar to the
South Pacific necessary for the development of C. fili-
corne ?

The surface temperatures at the stations where the
species occurred, varied from 19.°0 to 28.°7 C. The en-
vironmental conditions in situ were: temperature, 14.°3
to 27.°6 C; salinity, 34.4 to 36.3 per mille; pH, 7.17 to
8.27; phosphate, 8 to 32 mg P04/m3.

There was only one record from water with less than
10 mg P04/m3. Nielsen (1934) designated C. filicorne
as a genvilne shade species, and this is borne out by the
Carnegie data, as can be seen in table 25. It was found
predominantly in the 100-meter samples, and only a sin-
gle specimen was fovuid in the surface samples.

Table 25.

Records of occurrence of C. filicorne
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

Total

0.3

1

0.4

4

1.3

3

1.1

11

4.3

1

0.4

6

2.3

0.3

1.9

21

7.9

A=Number of records. B=Per cent of total number of
samples collected at that depth

30. Ceratium symmetricum Pavillard
Figures 15H-L, chart 24, appendix table 28

Paulsen (1931) suggested that the name C. gracile be
supplanted because of the confusion of synonyms and the

differences of opinion of Jorgensen (1911, 1920) and
Pavillard (1923). The present authors agree that the
name should be dropped, not because of the confusion of
these authors' opinions, however, but because the origi-
nal drawing of Gourret (1883, pi. I, fig. 1) is not identi-
fiable.

There is a considerable difference of opinion as to
what segregation of forms should be made within this
group. Peters (1934) listed only two forms; C. gracile
gracile and C. gracile svmmetricum. He included var.
orthoceros Jorgensen in his subspecies gracile. It is
doubtful, however, if he found coarctatum . since his draw-
ings do not show it. Nielsen (1934), on the basis of some
statistical work on size and horn spread, decided that the
various forms he found could be lumped into a single uni-
fied species. It is doubtful, however, if his measurements
of horn spread properly express the difference in body
shape and origins of horns--which are the distinguishing
features of the segregates of this group. The present au-
thors believe, on the basis of the study of the Carnegie
material, that coarctatum and orthoceros are sufficiently
distinct to justify the retention of their names and the
recording of their distributions. There is sufficient in-
tergradation between them, on the other hand, that they
cannot be given more than varietal value. Variety coarc-
tatum is characterized by long, more or less parallel,
horns; var. svmmetricum by widely spreading horns;
var. orthoceros by short close horns, and the right ant-
apical in particular, with an abrupt anterior turn.

The three forms of the species are not subspecies
in the geographic sense; they are all found within the
same regions. For this reason their records have been
combined on the chart (chart 24) although the individual
records are noted on the distributional list (appendix table
28).

The species as a whole is a fairly common warm-
water form. In the Carnegie collection it was found at
eighty-one stations- -fifteen in the Atlantic and sixty-six
in the Pacific. In the Atlantic these were all in the warm
Atlantic region. The surface temperatures at these sta-
tions were all above 24.°0 C. In the Pacific the species
occurred in the warm Pacific region and throughout the
southeast Pacific region. At the northern limits of the
warm Pacific region the species occurred where the sur-
face temperatures were as low as 19.°1 C. In the south-
east Pacific region It occurred at surface temperatures
as low as 15.°0 C. Thus, it was classified as a slightly
tolerant tropical species.

Table 26.

Records of occurrence of C . symmetricum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

10
2

2.6
0.5

31

23

3

11.0
8.1
1.6

45

34

8

17.7

13.4

3.2

Total

12

3.1

57

20.7

87

34.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

Its relative abundance is also shown by the number
of sample records. There were 156 recordsof occurrence,
of which 86 were rare, 59 occasional, and 11 common. It

30

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

was found more frequently in the net samples, with 100
net records and 56 pump records.

Ceratium svmmetricum is a pronounced shade spe-
cies. Nielsen (1934) usually found it in the samples col-
lected below the 50-meter level except at stations rich
in plankton. The Carnegie dat a indicate a much greater
frequency in the deeper levels (see table 26). Only 12 of
the 156 records, or 3.1 per cent of the surface samples,
were from the surface, whereas the species was found
in 34.0 per cent of the 100-meter samples.

The species is quite common in oligotrophic water.
Sixty -five of the 156 records of occur rence were in water
containing less than 10 mg P04/m3. The environmental
conditions in situ were: temperature, 12.°1 to 29.°3 C;
salinity, 30.7 to 37.1 per mille; pH, 7.90 to 8.39; phos-
phate, 3 to 152 mg P04/m3.

31. Ceratiimi axlale Kofoid
Figures 15D and E, chart 25, appendix table 29

This is a very rare intolerant tropical species. It
was found only twice in the collections of the Meteor in
the South AUantic (Peters, 1934). Nielsen (1934) found
it at nine stations in the Pacific in the South Equatorial
and East Australian currents. The Carnegie collections
extend the known range of the species both in the North-
ern and Southern hemispheres of the Pacific. It was not
found in the Atlantic. It occurred at eleven Carnegie sta-
tions; two of these were in the southeastern Pacific, one
off Japan, and the rest in the North Equatorial Current
and north and east of Hawaii. It is probable that the
species is more or less evenly distributed over the warm
waters of the world but it is seldom collected because of
its sparse numbers. In the Carnegie collection there
were thirteen records of occurrence, of which eight were
rare, three occasional, and two common.

The surface temperatures at the stations where it
occurred, varied from 20.°2 to 28.°1 C. The ranges of en-
vironmental conditions in situ were: temperature, 15.°6
to 27.°6 C; salinity, 34.5 to 35.1 per mille; pH, 8.08 to
8.39; phosphate, 4 to 32 mg P04/m3.

Five of the thirteen records were in water contain-
ing less than 10 mg P04/m3.

All of Nielsen's (1934) records were from below 50-
meter depths except one. The Carnegie data are in good
agreement with his (see table 27). There was only one
record from a surface sample and one from a 50-meter
sample. The other eleven were from 100-meter sam-
ples. It is thus evident that C. axiale is unquestionably
a shade species.

It is also of interest to note that eight of the records
were from pump samples, whereas only five were from

Table 27.

Records of occurrence of C. axiale
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

1

0.3

1 0.4

6 2.4
3 1.2

2 0.7

Total

1

0.3

1 0.4

11 4.3

A=Number of records. B=Per cent of total number of
samples collected at that depth.

the net samples. Apparently the curved nature of the
antapical horns permits the specimens to pass through
the coarser nets.

32. Ceratiimi aultii n. sp.
Figures 15F and G

Diameter, 55 microns; total length, 230 to 250 mi-
crons. The shape of the body is similar to those of C^
svmmetricum and C. axiale . except that the apical horn
is displaced to the left. Curvatures of antapical horns
resemble those of C. axiale except at the bases where
the antapicals extend laterally from the body before they
curve anteriorly.

Ceratium aultii was found at only two stations — one
off Peru (station 72) and the other in the central part of
the North Equatorial Current of the Pacific (station 151).
It was found at 100 meters at both stations. At station
72 it was "common" in the net sample; at station 151 it
was "rare" in the pump sample. The surface tempera-
tures at the stations where it was found were 24.°9 and
26.°0 C. The temperatures in situ were: 14.°8 at station
72, and 12.°5 C at station 151.

33. Ceratium azoricum Cleve
Figures 16M-P, chart 26, appendix table 30

This species is a variable and little-known form. It
is probably much more common than the number of rec-
ords for it would indicate, since it is lost through the net
because of its small size. It varies greatly in size (di-
ameter from 38 to 50 microns), however, as well as in
shape of body, and origin of antapical horns. The larger
forms with spreading horns bear a close resemblance to
C. peter sii and may, in fact, be identical with this, i.e.,
C. peter sii may represent simply one end of a line of
variations of C. azoricum . At the other end of this line
are the small specimens with horns close to the body.
In the Carnegie collection a specimen was found which
had a heavy list which joined the apical with the antapical
horns (fig. 16M). This may be a new species as no defi-
nite intergrades were found between it and C. azoricxmi .
Since only the one specimen was found, however, it was
decided to postpone naming a new species until more
material could be examined.

Ceratium azoricum is a very tolerant tropical spe-
cies. Lebour (1925) stated that it is subtropical and is
occasionally brought into the northern seas. In the Car-
negie collection it was found in the region of Ireland and
between Iceland and the Faeroes. In the latter region the
surface temperature was 8.°9 C. In the Pacific it was
found at one station in the cold North Pacific region,
where the surface temperature was 13.°3 C. In the south-
eastern Pacific it was found at temperatures as low as
16.°3 C. On the other hand, it occurred at temperatures
as high as 29.° 5 C in the equatorial regions. It was foimd
at a total of five stations in the Atlantic, only one of which
was in the warm Atlantic region. In the Pacific it was
found at twenty-three stations: one in the cold North Pa-
cific region, seven in the southeast- Pacific region, and
fifteen in the warm Pacific region. The total number of
records was sixty, of which thirty-eight were rare, twenty-
one occasional, and one common. As stated above, the
species is probably commoner than the reported records
would indicate because it probably escapes through the

CERATIUM SPECIES OF CARNEGIE COLLECTION

31

nets. Nevertheless, it was found more often in the net
than the piimp samples. There were thirty-three rec-
ords (16 per cent) for the net samples and twenty-seven
records (14 per cent) for the pump samples.

Nielsen (1934) stated that C. azoricum is a typical
surface species but this is not confirmed by the Carne-
gie data (see table 28). There are not sufficient deeper
records, however, to warrant classifying C. azoricum as
a typical shade species.

Table 28.

Records of occurrence of C. azoricum Cleve
at three levels

Occur -

Depth in meters

rence

50

100

A

B

A

B

A

B

Rare

Occasional

Common

16
6

1

4.2

1.6
0.3

14

8

5.0
2.8

8

7

3.2

2.8

Total

23

6.1

22

7.8

15

6.0

A=Number of records. B=Per cent of total number of
samples collected at that depth.

The ranges of environmental conditions in situ were:
temperature, 8.°2 to 29.°3 C; salinity, 32.7 to 36.0 per
mille; pH, 7.85 to 8.39; phosphate, 7 to 176 mg P04/m3.

Only two of the sixty records of occurrence were
from water containing less than 10 mg P04/m3.

34. Ceratium petersii Nielsen
Figure 16L, chart 26, appendix table 31

As noted above, it is questionable whether this form
should be considered separate from C. azoricum Cleve
(see p. 30). For the present, however, it will be con-
sidered separately. Nielsen (1934) reported the species
from southeast of New Zealand. The records of Peters
(1934) for C. bucephalimi in the South Atlantic were re-
ferred to C. petersii bv Nielsen (1934), but the present
authors cannot agree with this (see below). The Carne-
gle records for C. petersii were at six stations, all in
the Pacific--five in the southeastern Pacific and one off
the west coast of the United States. Thus it is probable
that C. petersii is peculiar to the Pacific Ocean.

Geographically it has been difficult to classify C.
petersii . partly because of the limited number of rec-
ords and partly because of Its peculiar distribution. It
is either cosmopolitan or subpolar. The surface tem-
peratures at its record stations varied from 16.°5 in the
South Pacific and 16.°4 in the North Pacific to 21. °2 off
Peru. It could be classified as subpolar except for its
peculiar distribution in the southeastern Pacific. No
other subpolar form has shown such a distribution. It
is tentatively considered to be a cosmopolitan species.
There is, of course, the possibility that C. petersii is a
cold-water subspecies of C. azoricum but this possibil-
ity is lessened by the fact that the typical C. azoricimi ls
also found in cold water.

The total number of sample records for C. petersii
in the Carnegie collection was fifteen, of which ten were
rare, four occasional, and one common. Eight of the
records were from net samples and seven were from
pump samples.

The ranges of environmental conditions in situ were:

temperature, 10.°8 to 21.°2 C; salinity, 33.0 to 35.1 per
mille; pH, 7.68 to 8.18; phosphate, 12to233mgP04/m3.

None of the records were in oligotrophic water.

Nielsen (1934) found. the species only in the samples
from 50 to meters. In the Carnegie collections there
were six records for the surface, six for 50 meters, and
three records for 100 meters. Thus it is probably a sur-
face species.

35. Ceratium arietinum Cleve
Figures 16A-K, chart 27, appendix table 32

This species is one of the most ill defined of the
genus. It is composed of a great many variants, includ-
ing C. hur.ephaliiTTi which is here treated as a subspecies.

Peters (1934) stated that in the Meteor material all
the variation forms described by Jorgensen (1911 and
1920) were foimd, but two forms were more common--a
small one and a large one. These intergraded but f. gracl-
lentum was distinct.

The "large and small main form C. arietinum arie-
tinum " was found in various regions of the South Atlantic
but f . gracilentum was purely tropical, occurring between
latitudes 12° north and 10° south with water temperatures
over 22° C.

Nielsen (1934) stated that he found only one form in
the South Pacific and that it corresponded to Jorgensen's
f. detortvun and f. gracilentum. Thus, apparently, Niel-
sen considered these two forms identical.

The Carnegie collection is particularly rich in vari-
ations of this species. Only the more important of these
have been figured (figs. 16A-K).

The characteristic arietinum has the apical horn dis-
placed to the left, whereas the right antapical horn is
strongly incurved (fig. 16E). In f. gracilentum Jorgensen
there is a distortion and lengthening of the apical horn,
which is sharply curved proximally (fig. 16K). Jorgensen
(1920) on the other hand, described a form, f. regulare .in
which the apical horn is symmetrically placed. This
form resembles C. bucephalum Cleve.

In the present collection the placement of the apical
horn was extremely variable and showed transitions to
C. bucephalum (see figs. 16A-H).

Nielsen (1934) classified Peters' (1934) southernform
of C. bucephalum as a new species, C. petersii. The
drawing of this species which Nielsen gives, however, is
quite different from the one given by Peters. In our opin-
ion these two forms cannot be included together. Peters'
figure (plate 3, fig. 16c) is truly a form of C. bucephalum .
which the authors have included as a subspecies of C.
arietinum. Nielsen's figure (fig. 44), however, is proba-
bly a distinct species as he indicated. It is closely re-
lated to C. azoricum (which see).

The gracilentum form of the Pacific (fig. 16F) is dif-
ferent from that of the Atlantic (fig. 16K). The body is
not nearly so distorted, the antapical contour is more
nearly perpendicular to the axis of the apical horn, the
body is relatively wide, and the apical horn is not so ex-
tended. The transitions between gracilentimi and the typ-
ical arietinum. however, are continuous in both oceans.
It is very probable that several subspecies (in the
geographic sense) are concerned in the present complex.
Peters (1934) found f. gracilentimi to be limited to the
purely tropical waters between latitudes 12° north and 10°
south with temperatures over 22° C. In the Carnegie col-
lection this form was found farther north (to latitude 38.° 5,

32

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

station 15) but the temperatures were quite high (2i°8 C).
In the Pacific this particular form does not occur. Its
representative, however, (fig. 16F) is a warm-water
form, although not markedly so because the form in the
Pacific is not so distinct.

Bucephalum is truly of temperate- and cold-water
origin. Figures 16D and E show the form from the North
Pacific region, figure 16A from the Humboldt Current,
and figure 16C from the southeast Pacific region. It is
absent from the tropics and thus is bizonal as Peters
(1934) first demonstrated.

Since bucephalum showsthis geographic distribution,
and since it Intergrades with C. arietinum . it should be
considered a subspecies of C- firiptinnm- Jorgensen's f.
gracilentum is also a subspecies since it is limited to
quite warm water. It is useless, however, to name the
many variations and intergrades which occur among the
subspecies as they are innumerable and apparently of no
ecologic significance.

The species as a whole is a fairly common one. In
the Carnegie collection it occurred at sijrty-two stations
--ten in the Atlantic and fifty-two in the Pacific. The
total number of records was 123, of which 87 were rare,
30 occasional, 4 common, and 2 abundant. There were
67 net records and 56 pump records.

Nielsen reported C. arietinum as an outstanding
deep-water species. This is substantiated by the Car-
negie observations, which show the species much more
frequent at the 100-meter level than at 50 meters or the
surface (table 29). It was least frequent at the surface.

Table 29.

Records of occurrence of C.
at three levels

arietinum

Occur-

Depth in meters

rence

50

100

A B

A B

A

B

Rare

23

5.6

24

8.5

40

15.8

Occasional

6

1.6

13

4.6

11

4.3

Common

1

0.3

2

0.7

1

0.4

Abundant

1

0.3

1

0.4

Total

31

7.8

40

14.2

52

20.5

A=Number of records. B=Per cent of total number of
samples collected at that depth.

As indicated by the wide range of the species, it is a
rather eurythermal form. In the Atlantic it occurred at
surface temperatures ranging from 11° to 27.°5 C. In the
Pacific the range was greater; from 6.°9 (northeast of
Japan) to 29.°5 C (in the central tropics). The total ranges
of environmental conditions in situ were: temperature,
l.°6 to 29f5 C; salinity, 32.7 to 36.5 per mille, pH, 7.85
to 8.39; phosphate, 3 to 193 mg P04/m3.

The species was found more frequently in the richer
waters. Only 17 of the 123 records of occurrence were
from water containing less than 10 mg P04/m3.

36. Ceratium decluiatimi Karsten
Figures 16Q-T, 17A-C, chart 28, appendix table 33

This is a very variable, slightly tolerant tropical
species. Jorgensen (1911, 1920) made several varieties
among the variants of the species. Peters (1934) desig-
nated three subspecies in his South Atlantic material;
C. declinatum declinatum . C. declinatum angusticornum .

and C. declinatum laticornum. Melsen (1934) stated that
he could not distinguish between Peters' angusticornum
and laticornum . and in his South Pacific material he rec-
ognized only two "subspecies" — declinatum and angusti -
cornum. These two forms occurred in the same regions
so they are not subspecies in the sense used by Peters,
who foimd a certain geographic segregation of the vari-
ants. When a world collection is studied, such as that of
the Carnegie , it becomes evident that the species con-
sists of a large number of intergrading units. The nam-
ing of these serves no useful purpose. They are not
subspecies so far as can be determined, since they oc-
cur indiscriminately throughout the entire range of the
species.

Seven of these variants are shown in figures 16Q-T
and 17A-C. Some of these can be identified with named
forms of other authors. Thus, figure 16S is similar to
Jorgensen's var. maius: figure 16T to Peters' laticornum:
figure 16Q to declinatimi : figures 17A to C to angusti-
cornum: whereas figure 16R, with outturned right antap-
ical, has not been named.

In the Carnegie collection the species as a whole was
found rather continuously throughout the warm Atlantic,
warm Pacific, and southeast Pacific regions. It occurred
at a total of ninety-eight stations- -fourteen in the Atlan-
tic and eighty-fovir in the Pacific. In the Atlantic it oc-
curred at twelve stations in the warm Atlantic region and
at one in the cold Atlantic region, where it doubtless was
displaced by the Gulf Stream (station 3). In the Pacific
it did not occur in the cold North Pacific region but was
found off California at temperatures of 19.°1 and 16°2 C
at the northern part of the warm Pacific region. It oc-
curred in the southern part of the southeast Pacific re-
gion at stations where the surface temperatures were as
low as 16.°9 C. In the warm Pacific region it occurred
where the surface temperatures were as high as 29.° 5 C.

The environmental conditions in situ were as follows:
temperature, 13.°6 to 29.°4 C; salinity, 31.7 to 37.1 per
mille; pH, 7.17 to 8.42; phosphate, 3 to 123 mg P04/m3.

The ubiquity of this species (within the tropics) is
shown by the number of records, which was 204, of which
133 were rare, 67 occasional, and 4 common. Since it is
a very small species, it was found more commonly in the
pump samples. It occurred in 87 (16 per cent) net sam-
ples and 117 (32 per cent) pimip samples.

It seemed to show no preference for oligatrophic
water. Seventy-six of the 204 recordsof occurrence were
from water containing less than 10 mg P04/m3.

Nielsen (1934) stated that C. declinatum is definitely
a surface form. The Carnegie records show a decrease
in frequency with increase in depth (see table 30), but by
no means do they indicate that the species is exclusively

Table 30.

Records of occurrence of C. declinatum
at three levels

Occur-

Depth

in meters

rence

50

100

A

1 B

A

1 B

A B

Rare

Occasional

Common

60

34

3

15.6
8.8
0.8

36

19

12.7
6.7

37 14.5

14 5.5

1 0.4

Total

97

25.2

55

19.4

52 20.4

A=Number of records. B=Fer cent of total number of
samples collected at that depth.

CERATIUM SPECIES OF CARNEGIE COLLECTION

33

a surface form even when due correction is made for
contamination of the deeper net hauls. For Instance, it
was found in 25 per cent of the surface samples, but also
In 21 per cent of the 100-meter samples.

37. Ceratium gibberimi Gourret
Figures 17D-G, chart 29, appendix table 34

This Is a common, slightly tolerant tropical species
which occurs rather uniformly throughout the warm
waters of the world. Two varieties or subspecies are
usually recognized. One.f. dispar Pouchet (fig. 17D),
has a strongly curved right antapical horn and is com-
mon, whereas the other, f. subaeguale jSrgensen (fig.
17F), has regularly curved horns and is rare. The latter
was found by Peters (1934) in the South Atlantic in dif-
ferent regions, but it has not been reported before for
the Pacific. In the Carnegie collections one specimen of
this form was found (fig. 17F). This occurred off Japan at
station 114; depth, 50 meters; temperahireinsitu, 16.°2C.

In f . dispar there is considerable variation in the
height of the epltheca, curvature of horn, and placement
of apical horn, but no further segregation of the group
could be made. The Pacific and Atlantic specimens ap-
peared to be identical. A two-celled chain was found at
station 48 at 100 meters (fig. 17E).

The species was foiind at a total of 115 stations- -23
in the Atlantic and 92 in the Pacific. It was foimd at all
but three stations in the warm Atlantic region, but not
outside this region in the Atlantic (chart 29). In the Pa-
cific it was found almost continuously at the stations in
the warm Pacific region. It also occurred at ten stations
in the southeast Pacific region, but was absent from the
cold North Pacific region.

The total number of sample records for the species
was 297, with 179 rare, 105 occasional, and 13 common.
Of the total number, 221 were net records and 76 were
pump records.

The environmental conditions in situ were: temper-
ature, 14.°0 to 29.°3 C; salinity, 29.7 to 37.0 per mille;
pH, 7.17 to 8.47; phosphate, 2 to 178 mg P04/m3.

Nielsen (1934) found that C. gibberimi had its main
vertical distribution above 50 meters. The Carneede
data show that the species occ\u"s more frequently in the
upper layers, with a decrease in frequency down to 100
meters (table 31).

Table 31. Records of occurrence of C. gibberum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

Total

75

58

7

19.4

15.0

1.8

56

25

4

20.0
8.9
1.4

48

22

2

8.7
0.8

140

36.2 85 30.3 72 28.4

A=Number of records. B=Per cent of total number of
samples collected at that depth.

38. Ceratium concillans Jorgensen
Figures 17H-I, chart 30, appendix table 35

This species was formerly included in C. gibberum
ijut was separated by Jorgensen (1920) and this separation
has been accepted by later workers (Peters, 1934; Niel-

sen, 1934). Bohm (1931), however, figured C. gibberum
under the caption of C. concillans Jorgensen.

Peters (1934) considered this species to be an indi-
cator of warm oligotrophic water in the South Atlantic.
Nielsen (1934), however, found that in the South Pacific
the species is found as much in eutrophlc oceanic water
as in oligotrophic oceanic water. It did, however, seem
to avoid neritic conditions. In the Carnegie collection in
the Pacific it was found in both eutrophlc and oligotroph-
ic water. This feature could not be considered for the
Atlantic as it was found at only one station there (in oli-
gotrophic water). On the Carnegie cnxise it was found in
eutrophlc water in the southeastern Pacific, again off
Peru, and off California. On the other hand, it occurred
in many waters poor in plankton and nutrients. Thus,
thirty of the seventy-five records of occurrence for the
species were in water containing less than 10 mg
P04/m3. It must be concluded that C. concilians is not
limited to one type of water.

The species is not very common. In the Carnegie
collection it was found at a total of forty-two stations,
only one of which was in the Atlantic. The number of
sample records was seventy-five, of which fifty-nine
were rare, fourteen occasional, and two common. It is
much less common than its close relative, C. gibberum .
Since it is a fairly large form, it was found more fre-
quently in the net samples. There were forty-seven net
records and twenty-eight pump records.

The range of surface temperatures for the species
is rather large for a tropical form, namely, from 16.°4
(station 128) to 29.°4 C (station 95). The ranges of en-
vironmental conditions in situ were: temperature, 12.°1
to 29.°3 C; salinity, 33.0 to 36.4 per mille; pH, 7.71 to
8.47; phosphate, ,3 to 198 mg P04/m3.

Nielsen (1934) designated C. concilians as a surface
form. According to the Carnegie data, however, it is
found about equally at the surface, 50 meters, and 100
meters (see table 32).

Table 32. Records of occurrence of C. concilians
at three levels

Occ»ir-

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

Total

24

5.3

17

6.0

18

7.1

8

2.0

4

1.4

2

0.8

2

0.5

34

7.8

21

7.4

20

7.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

A form was found at station 130 (fig. 171) which has
rather regular antapical horns and thus represents a va-
riety of C. concllans corresponding to var. subaeguale of
C. gibberum .

39. Ceratium lunula Schlmper
Figures 17J-N, chart 31, appendix table 36

Two forms of this species are commonly recognized;
f . brachvceros with short apical horn and comparatively
long antapicals; and f. magaceros with somewhat shorter
antapicals but with long apical horn. The latter form

34

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

represents the anterior cell of a chain, whereas f . brachv -
ceros represents the remaining cells in the chain. It is
inconsistent to name the two elements of a chain in this
species alone as similar differences occur in the anteri-
or and following cells of the other species as well.

In the Atlantic the species is distributed over the
warm Atlantic region, extending to latitude 39° north,
but it is absent from the cold North Atlantic region.

The distribution of C. lunula at Carnegie stations in
the Pacific is unique among the Ceratium species (chart
31). It is similar to that of C. breve but even more re-
stricted (see p. 27). It is definitely a tropical species
and its distribution within the limits of 17° south and 20°
north latitude is fairly continuous. It was not found out-
side these limits except In the western Pacific, where it
extended in a continuous line to about 38 ° north off Japan
(station 115). It was conspicuously absent in the loop of
stations between the United States and Hawaii. It was
also notably absent from all the stations in the south-
eastern Pacific south of 20° south. An explanation for
this peculiar distribution is difficult to find. Certainly
it has no relation to temperature as many of the stations
where it does not occur have much higher temperatures
than many of the stations where it was found. The nutri-
ent content of the water cannot be a determining factor
either. The area north and east of the Hawaiian Islands
is extremely oligotrophic but so also is the region be-
tween Japan and Guam in the western Pacific, where the
species was foimd in large nimibers. Conversely, many
stations in the southeast Pacific region are in eutrophlc
water and yet did not support this species. Is it possible
that this species normally is confined within 20° of the
equator in the Pacific except off Japan where it is dis-
placed by the Kuroshio, or are the Carnegie records not
representative?

Peters (1934) did not find C. lunula south of 30° south
latitude in the South Atlantic. The records of the Dana
(Nielsen, 1934) for the South Pacific fit in well with the
distribution compiled from Carnegie data. Most of the
Dana records duplicate records of the Carnegie between
Panama and Samoa but its records extend somewhat west
of Samoa. Nielsen, however, did not find the species
south of 20° south. Thus, It is probable that the Carne-
gie records are representative and that the species nor-
mally is as restricted as these records indicate. Since
the species Is displaced northward off Japan, it is sur-
prising that it is not also displaced southward in the
region of Easter Island.

The total number of station records for the species
In the Carnegie collection was sixty-eight, of which twelve
were from the Atlantic and fifty-six from the Pacific. The
total number of sample records was 144, of which 88 were
rare, 52 occasional, and 4 common. The species was
found much more frequently In the net samples, with 121
net records and only 23 pump records. Thirty of the 144
records were in water containing less than 10mgPO4/m3.

Ceratium lunula was always foimd in water of high
temperature. In the Atlantic the surface temperatures at
its stations were all above 24° C, except one which was
20.°5 C. In the Pacific the surface temperatures were
above 20° except at one station off Japan, where it was
19.°9 and at the Galapagos where it was 18.°7 C. The
ranges of environmental conditions in situ were: temper-
ature, ll.°4 to 29.°3 C; salinity, 29.7 to 37.0 per mllle;
pH, 7.71 to 8.47; phosphate 3 to 220 mg P04/m3.

Ceratium' limula Is probably a shade species which
comes to the surface in waters rich in plankton. Nielsen

(1934) found it in the deeper hatils in the stations poor in
plankton. Jorgensen (1920) found it at greater depths in
the Mediterranean during the summer months. The Car-
negie records, not segregated according to richness of
plankton, show the highest percentage of records at 50
meters and a greater percentage at 100 meters than at
the surface (table 33). These records thus corroborate
the classification of this species as a shade form.

Table 33.

Records of occurrence of C. lunula
at three levels

Depth

in meters

rence

50

100

A

B

A

B

A

B

Rare

Occasional

Common

35

15

3

9.1
3.9
0.8

30
24

1G.7
8.5

23 9.0

13 5.2

1 0.4

Total

53

13.8

54

19.2

37 14.6

A=Number of records. B=Per cent of total number of
samples collected at that depth.

40. Ceratium carnegiei n.
Figures 18A-C

sp.

Transdiameter 92.5 (90 to 100) microns (six speci-
mens measured). Curvature of antapical horns similar
to that in C. lunula . Apical horn long and straight or
slightly curved to right. Body of epitheca slightly to
greatly inflated. In extreme cases the walls are parallel
for about one transdiameter from the girdle, at which
point they may be expanded to greater than girdle diam-
eter. Body walls thick and covered with numerous lists
as In the heavy specimens of C. lunula .

The species was foimd at four stations, all in the
Pacific. At one of these, station 71 off Peru, the body
was not much inflated. The most extreme cases were
found at the warmer stations, stations 151, 152, and 158.
It occurred at the surface at station 71, but at 50- and
100-meter depths at the other stations. The surface tem-
peratures at the stations where it was found, varied from
23.°5 to 28.°2C. The environmental conditions in situ were:
temperature, 12.°5 to 27.°6C; salinity, 35.9 to 34.0 per
mille; pH, 7.87 to 8.39; phosphate, 48 to 58 mg P04/m3.

41. Ceratium contortum Cleve
Figures 18D-N, chart 32, appendix table 37

There has been much confusion in the literal re-
garding of the taxonomy of both C. contortum Cleve and
C. karstenli Pavlllard ( C. arcuatum Cleve). After a
thorough study of the extensive material of the Carnegie
collection, the writers came to the conclusion that these
two species represent variants of a single species. The
difficulties of previous workers probably have been owing
in large part to the fact that they were attempting to dif-
ferentiate between these two forms when in fact they do
not exist and the separation is an artifical one.

Thus, Jorgensen (1911) In his monograph gives C.
karstenli . C. longlnum . and C. contortum as distinct spe-
cies. Ceratium contortvun is characterized by slightly
distorted body and slightly inward-bent right antapical
horn, except in var. saltans Schroder, which has a v'ry

CERATIUM SPECIES OF CARNEGIE COLLECTION

35

decidedly distorted body and sharply bent right antapical
horn. Jorgensen's figure of C. karstenii (fig. 117a),
however, shows a specimen with a curled right antapical
horn, which is certainly to be considered a double bend-
ing and represents a zone of instability in the same po-
sition as the bending In the horn of contortum. He rep-
resented C. longinum as more slender than C. karstenii
and with longer and straight antapical horns. He also
separated from C. karstenii . var. robustum . a variety
which is more robust than the main species and has
shorter and straight antapical horns. This form resem-
bles C. tripos.

Later, Jorgensen (1920) foimd all intergrades between
C. karstenii and var. robustimi so he no longer consid-
ered it even a good variety.

Peters (1934) treated C. longinum as a variety of C.
karstenii. He apparently considered saltans a distinct
variety of C. contortum. but does not give a figure of the
mainspecies. His figures of C. karstenii and C. contortum
(figs. lOe and g, plate 2) show equally distorted bodies.
Nielsen (1934), in his study of the Dana South Pacific
material, treated C. karstenii and C. contortimi sepa-
rately. He agreed with Peters (1934), however, thatvar-
robustum cannot be separated from C. karstenii. He
stated that he did not find C. longinum in his collections
but his figure of C. contortum f . subcontortum (fig. 52)
answers well to this form. He considered it a shade
form of C. contorhmi.

In the Carnegie collections every possible intergrade
was found between C. contortum . C. lohginum . and C.
karstenii . and the varieties robustum and saltans (see
fig. 18). There are heavy forms and light forms; some
with long horns and some with short. There are speci-
mens with kinked horns, but not much distorted body;
and, on the other hand, specimens with regular body, but
bent horns. The right antapical may bend in or out, or
may trail off in an attenuated manner. The extreme
forms were represented by var. saltans and usually were
smaller than the others and it was thought for a wMle
that a separation of two species could be made on this
basis. After measiiring many specimens from all local-
ities, however, it was foimd that this was impossible.
There is indeed a great variation in size. The transdl-
ameters range from 52 microns in the smaller specimens
of saltans to 110 microns in the more robust specimens.
There was no segregation into size groups, however; the
greatest number of specimens had transdiameters rang-
ing between 70 and 85 microns.

Ceratium contortum is a common, slightly tolerant
tropical species. It was found at practically all stations
in the warm Atlantic, warm Pacific, and southeast Pa-
cific regions, but was completely absent from the cold
North Atlantic and cold Pacific regions (chart 32).

Nielsen (1934) listed C. longinum Karsten as an At-
lantic form absent from the Pacific. It occurred in the
Carnegie collection, however, at many stations in the
Pacific.

The commonness of the species is shown by the
large nimiber of sample records, namely, 461, of which
194 were rare, 178 occasional, 84 common, and 5 abun-
dant. The net records for the species totaled 327; the
pump records, 134.

The range of surface temperatures at the stations
where the species occiirred, varied from 16.°3 to 29.°5C.
The ranges of environmental conditions in situ were:
temperature, ll.°4 to 29.°4 C; salinity, 30.0 to 37.1 per
mille; pH, 7.17 to 8.47; phosphate, 2 to 138mgP04/m3.

Many of the specimens have attenuated and sometimes
curved antapical horns which are extremely thin-walled.
It is probable that this is an adjustment to shade condi-
tions. Nielsen (1934) considered the subcontortum form
(longinum ?) a shade form of C. contortum. In the Car-
negie material var. saltans showed a greater frequency
at greater depths than did the more regularly-shaped
forms. Although the more robust forms sometimes oc-
curred with thin antapicals, this condition was much
more common in the saltans group. In the others the
larger specimens were much more predominantly sur-
face forms. Among the specimens more than 80 microns
in diameter, there were twice as many surface records
as 50-meter records and very few at 100 meters. Of the
specimens less than 80 microns in diameter, the number
of records at the siirface and 50 meters was about equal,
although the number at 100 meters was almost negligible.
Among saltans , too, the greatest frequency was at the
surface. The percentages for the species as a whole
show a predominance at the surface (see table 34).

Table 34.

Records of occurrence of C. contortttm
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional
Common
Abundant

Total

95

80

34

4

27.7

20.7

8.8

0.2

54
53
33

19.2
18.7
11.7

45 17.7

45 17.7

17 6.7

1 0.4

213

57.4 140 49.6 108 42.5

A=Number of records. B=Per cent of total number of
samples collected at that depth.

42. Ceratium limulus Gourret
Figure 19A, chart 33, appendix table 38

This is a rather rare intolerant tropical species
which Is quite characteristic and not very variable.
Peters (1934) considered this species an indicator of
warm ollgotrophic water in the South Atlantic; but Niel-
sen (1934) foimd it in both oUgotrophic and eutrophic
water in the South Pacific, with its greatest abundance in
eutrophic water. In the Carnegie collection it was found
only in ollgotrophic water in the Atlantic, but in both eu-
trophic and oUgotrophic water in the Pacific. Is it pos-
sible that the Atlantic form is less adaptable in this re-
gard than the Pacific form?

The species was found at fifty-five stations — nine in
the Atlantic and forty-six in the Pacific. The Atlantic
stations were confined to the warm Atlantic region where
the surface temperatures were high (from 24.°8 to 27.°6
C) and the nutrient content of the water was low. The
phosphate values for the upper 50 meters were all below
11 mg P04/m3. In the Pacific the stations were confined
to the warm Pacific region for the most part, although
there were six stations in the southeast Pacific region.
The range of conditions at the Pacific stations was much
greater than those at the Atlantic stations. For instance,
the surface temperatures varied from 18.°7 to 29.°3 C,
and the mean phosphate value for the upper 50 meters
varied from 4 to 83 mg P04/m3. It should be noted, how-
ever, that all stations were remote from continents.

36

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

The ranges of environmental conditions in situ for
the species at all stations were as follows: temperature,
13.°8 to 28.°2 C; salinity, 33.9 to 36 per mille; pH, 8.47
to 7.80; phosphate, 4 to 150 mg P04/m3.

Thirty of the eighty-eight records were from water
containing less than 10 mg P04/m3.

The total number of sample records for the species
was eighty-eight, of which eighty-five were rare and
three occasional. There was a large proportion of pump
records owing to the small size of the species (thirty-
nine pump records and forty-nine net records).

Ceratium limulus apparently lives at various depths
but is found somewhat more frequently in the upper lev-
els. Bohm (1931) found it in the Adriatic as deep as 200
meters. Nielsen (1934) found it predominantly in the 50-
to 0-meter samples. In the Carnegie collection the spe-
cies was found with decreasing frequency with increase
in depth to 100 meters (table 35), although the frequency
at 100 meters (in percentage) was more than half that at
the surface.

Table 35.

Records of occurrence of C.
at three levels

limulus

Occur -

Depth

in meters

50

100

A

B

A

B

A B

Rare
Occasional

39
2

10.1
0.6

28

1

9.9
0.4

18 7.1

Total

41

10.7

29

10.3

18 7.1

A=Number of records. B=Per cent of total number of
samples collected at that depth.

43. Ceratium paradoxides Cleve
Figure 19B, chart 34, appendix table 39

This is a rare Intolerant tropical species. Peters
(1934) found it in both oligotrophic and eutrophlc water
in the South Atlantic. Nielsen (1934) found it scattered
over the region traversed by the Dana from Panama to
Australia. It is never common and its distributional rec-
ords are discontinuous for this reason.

In the Carnegie collection the species was found at
sixty-four stations- -thirteen in the Atlantic and fifty-one
in the Pacific. In the Atlantic the stations were confined
to the warm Atlantic region. In the Pacific they were
restricted to the warm Pacific region and the southeast
Pacific region (see chart 34).

There were ninety-one records of occurrence, of
which eighty were rare and eleven occasional. Sixty-six
of these records were from net samples and twenty-five
were from pump samples.

Nielsen (1934) stated that C. paradoxides is a shade
species, and this is corroborated by the Carnegie obser-
vations. In fact, its vertical distribution is most decid-
edly stratified. As shown in table 36, there were only
six records for the surface samples (representing only
1.6 per cent of the samples), whereas the species was
found in 22.8 per cent of the 100-meter samples.

The surface temperatures at the stations where C.
paradoxides occurred In the Atlantic varied from 20.° 5 to
27.°6 C; in the Pacific from 19° to 29.°5 C. The environ-
mental conditions in situ were as follows: temperature,
12.°1 to 29.°3 C; salinity, 33.4 to 37.0 permUle; pH, 7.17

to 8.39; phosphate, 4 to 123 mg P04/m3. It was found
in both eutrophlc and oligotrophic water. Thirty -four of
the ninety-one records were in water containing less than
10 mg P04/m3.

Table 36. Records of occurrence of C. paradoxides
at three levels

Occur-

Depth in meters

50

100

A

1 B

A B

A B

Rare
Occasional

5

1

1.3
0.3

26 9.2 49 19.3
1 0.4 9 3.5

Total

6

1.6

27 9.6 58 22.8

A=Number of records. B=Per cent of total number of
samples collected at that depth.

44. Ceratium platycorne Daday
Figures 19C-H, chart 35, appendix table 40

This is a rare, slightly tolerant tropical species.
Varieties of the species have been described, but these
are of no particular importance. Variety cuneatxmi Jor-
gensen is not distinguishable from the main species. Its
shorter truncated antaplcal horns probably are owing to
autotomy or accidental breaking of the distal part of the
horns. It is probable that the jagged edges of these
horns never represent the original condition. Figure
19D shows a specimen with broken (truncate) right ant-
aplcal which is shorter than the left. The right is nor-
mally longer than the left. Variety lamelllccrne is some-
what more distinct. Nielsen (1934) found it quite distinct
in the Dana collections. It is, however, simply a narrow-
horned variant and Intergrades are not absent (fig. 19G).

Ceratium platycorne is widespread over the tropi-
cal waters of the world and is sometimes displaced by
currents. It has been found by others in the northern
waters of the Atlantic (Paulsen, 1908; Lebour, 1925). In
the Carnegie collection it occurred off Iceland, with a
surface temperature of 12.°44 C. Otherwise, in the At-
lantic it was confined to the warm Atlantic region, with
surface temperatures above 20° C. In the Pacific it was
limited to the warm Pacific and southeast Pacific re-
gions, with surface temperatures ranging from 19.°27 to
28.°74 C. The ranges of environmental conditions in situ
were: temperature, 12.°4 to 27.°6 C; salinity, 34.1 to
37.0 per mille; pH, 7.87 to 8.39; phosphate, 4 to 152
mg P04/m3.

Peters (1934) found C. platycorne widespread in
oligotrophic water in the South Atlantic, and Nielsen
(1934) found it in the entire region between Panama and
Australia. In the Carnegie collection it did not show any
preference for one type of water. Eighteen of the fifty-
three records of occurrence were from water containing
less than 10 mg P04/m3.

The species is not common. There were onlyforty-
tliree stations at which It was found, of which fourteen
were in the Atlantic and twenty-nine in the Pacific. The
total number of records (fifty-three) is relatively small.
Of these, forty-four were rare, eight occasional, and
one common. There were thirty-eight net records, and
fifteen pump records.

Ceratium platycorne is distinctly a shade species.
The records indicate this even more definitely than in

CERATIUM SPECIES OF CARNEGIE COLLECTION

37

the case of C. paradoxides. Of the fifty-three records,
only one was from the surface, whereas there were thir-
teen from 50 meters, and thirty-nine from 100 meters.
The percent.ages are even more striking (table 37).

Table 37. Records of occurrence of C. platycorne
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

11

3.9

32

12.6

2

0.7

6

2.4

....

1

0.4

Rare 1 0.3

Occasional

Common .... ^

Total 1 0.3 13 4.6 39 15.4

A=Number of records. B=Fer cent of total number of
samples collected at that depth.

45. Ceratium ranlpes Cleve
Figures 19I-K, 20, 21A, chart 36, appendix table 41

This is a rather common, slightly tolerant tropical
species. It is variable in respect to the number of "fin-
gers;" there may be three or four (fig. 21 A) or 19 and 21
(fig. 20). The "few-fingered" forms were formerly seg-
regated as var. ranipes of C.palmatum but, since a com-
plete intergradation between it and the main species is
commonly found, the variety recently has been disregard-
ed (Peters, 1934; Nielsen, 1934). There is a great varia-
tion in the length of theantapical horns. The long horns
usually have fewer fingers (cf. figs. 191, 21A). A specimen
was found in the Carnegie collection with an extremely
large number of fingers (fig. 20). There were nineteen on
the left horn and twenty-one on the right. Accordlngto the
theory of shade forms (Nielsen, 1934), extreme flattening
of the ends of the hind horns and their division into a num-
ber of linear branches represents an adaptation of the
species to shade conditions. This is substantiated by the
tact that the hind horns and fingers are densely packed
with chromatophores, and is conclusively shown by the
vertical distribution of species records (see below).

The horizontal distribution of C. ranipes is typical
of slightly tolerant tropical species. It shows a slight
tendency to drift out of the tropical regions. It was found
at a total of seventy-nine Carne^ e stations — twenty-one
in the Atlantic and fifty-eight in the Pacific. In the At-
lantic the stations were all confined to the warm Atlantic
region, with the exception of one (station 3) where the
species was displaced into colder water (surface temper-
ature 15.°5 C). In the Pacific the stations were llmitedto
the warm Pacific region and the southeast Pacific region.
The range of surface temperatures at the Pacific record
staUons was 18.°7 to 29.°5 C.

The ranges of environmental conditions in situ for
both oceans were as follows: temperature, 13.°1 to 29.°3
C; salinity, 33.7 to 36.8 permille; pH, 8.00 to 8.47; phos-
phate, 2 to 99 mg P04/m3.

Ceratium ranipes probably is more or less uniform-
ly distributed in all tropical waters. Peters (1934) foimd
a slight Indication of preference for oligotrophlc water in
the South Atlantic, but Nielsen (1934) found the species in
both eutrophlc and oligotrophlc water and even in neritic
conditions in the South Pacific. In the Carnegie collec-
tion no preference for oligotrophlc water was indicated.

Forty-seven of the 147 records of occurrence were in
water containing less than 10 mg P04/m3.

Ceratium ranipes is a fairly common species, as
indicated by Carnegie records. There was a total of 148
records of occurrence, with 116 rare and 32 occasional.
Thirty-nine of these were pump records and 108 were net
records.

Cerativmi ranipes is definitely a shade species. Niel-
sen (1934) found it at deeper levels except at stations rich
in plankton where it was found mostly above 50 meters.
In the Carnegie collection the species was found with in-
creasing frequency with increase in depth down to 100
meters. Expressed in percentage, the number of records
at 100 meters was more than twice that at the surface
(see table 38).

Table 38.

Records of occurrence of C.
at three levels

ranipes

Occur-
rence

Depth in meters

50

100

A

B

A B

A B

Rare
Occasional

30
8

7.8
2.1

42 14.9 44 16.9
12 4.2 12 4.7

Total

38

9.9

54 19.1 56 21.6

A=Number of records. B=Per cent of total number of
samples collected at that depth.

46. Ceratium macroceros (Ehrenberg) Vanhoffen
Figures 21B-F, chart 37, appendix table 42

Ceratium macroceros is a semicosmopolitan spe-
cies with a peculiar distribution. In the Atlantic there
are two subspecies (Jorgensen, 1911, 1920). One, com-
monly called the main species, is northern and extends
south to the Bay of Biscay and the northern parts of the
GuU Stream. For convenience, it is here named subsp.
macroceros n. subsp. The tropical form is subsp. galli-
cum Kofoid. Jorgensen (1920) found only subsp. gallicum
in the Mediterranean. Peters (1934) stated that in the
South Atlantic subsp. gallicum predominated everywhere
except in regions VI and VII (south of Africa and west-
ward in the general latitude of 40° south), where there
was a form similar to the main species of the North At-
lantic. Unfortunately, he did not figure this form. It may
be an antarctic subspecies Indicating a bipolarity in the
Atlantic. Nielsen (1934) foimd only subsp. gallicum in
the South Pacific. In the Carnegie collections in the At-
lantic the two subspecies characteristic of the North At-
lantic were found. Subspecies macroceros occurred
north of 40° north and at two stations south of this lati-
tude, stations 15 and 16. Conversely, subsp. g allicum
occurred principally at the tropical stations but also at
one station north of 40° north, station 13. The specimens
there, however, were not typical of the tropical subspe-
cies but were in the nature of intergrades.

The northern subspecies, subsp. macroceros n. subsp.,
differs from subsp. gallicum in its size, shape, and cur-
vature of antapical horns. The transdiameter of the north-
ern form is usually from 50 to 55 microns, whereas in
subsp. gallicum. it is less than 50 microns. The northern
form is more robust generally, and the body is somewhat
longer. The antapical horns in the two forms leave the
body at about the same angles but in subsp. gallicum they

38

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

are much more abruptly turned anteriorly and all their
horns are thinner (cf. figs. 21E and F).

The species as a whole is one of the most common
of the genus. It was found at 148 stations--42 in the At-
lantic and 106 in the Pacific. It was found frequently in
all the samples from a station so that the total number
of sample records for the species is large, namely, 482.
Of these, 226 were rare, 201 occasional, 54 common,
and 1 abundant. Net records predominated, of which
there were 307 as compared with 175 pump records.

When dealing with a species as common as this one,
a series of negative records is significant. In this con-
nection it is Interesting to turn to the distribution of the
species in the North Pacific (chart 37). Subspecies gal-
licum was found at every station in the warm Pacific re-
gion except one off Japan, where its northern distribution
in the west ended abruptly at about 38° north latitude. In
the east, off California, the northern limit was equally
abrupt but there it was above 40° north latitude, with one
station in the cold North Pacific region. Otherwise, no
representative of C. macroceros was found in the cold
North Pacific region. The absence of subsp. macroceros
in this area is a good indication of the planktologlc iso-
lation of the North Pacific (see p. 11).

In the South Pacific in this species we find evidence
of the continuity of the colder waters of all the southern
oceans. In the extreme southeastern part of the area
traversed by the Carnegie the species assumed the as-
pect of subsp. macroceros (fig. 21C). This is possibly
the southern form mentioned by Peters (1934) from the
Atlantic.

Subspecies g alllcum may be classified as slightly
tolerant tropical . Off Japan it extended into water with
surface temperature of 19.°9; off the west coast of Amer
ica to temperature of 16.°4, and in the southeast Pacific
to 15.°0 C. In the Atlantic subsp. galllcum occurred at
surface temperatures of about 20° C, except at station
13, where the temperature was 11. °3 C.

The ranges of environmental conditions in situ were:
temperature, -l.°6 to (29.°4) C; salinity, 31.7 to 37.1 per
mUle; pH, 7.17 to 8.47; phosphate, 2 to 123 mgP04/m3.

It should be noted that the species was not found at a
line of stations off Peru nor between the Galapagos Is-
lands and Panama with the exception of two stations. The
waters here, especially off Peru, are eutrophic; the phos-
phate content in the upper 50 meters is mostly above 50
mg P04/m3. Is it possible that this accounts for the ab-
sence of C. macroceros? The temperatures at these sta-
tions were mostly above 20° C, so temperature hardly
can be responsible. Possibly C. macroceros avoids high
concentrations of nutrients. In this connection it should
be noted that in the North Atlantic, where C. macroceros
occurs at low temperatures, the phosphate content in the
upper 50 meters is mostly below 50 mg, whereas in the
northern North Pacific, where the species is absent, it
ts mostly above 100 mg. These conditions must be taken
Into account in any attempt to explain the distribution of
this species. It was pointed out in the general discussion
(p. 7), however, that it Is not likely that the concentra-
tion of such plant nutrients as phosphate and nitrate found
In the oceans are important in the qualitative distribution
of Ceratlum species.

Ceratium macroceros occurs rather uniformly down
to the 100-meter depth. Nielsen (1934) found it mostly
above SOmeters, but in the Carnegie collection the great-
est percentage was at 50 meters, with slightly lower per-
centages at the surface and 100 meters (table 39).

Table 39. Records of occurrence of C. macroceros
at three levels

Omir-

Depth

in meters

rsncG

50

100

A

1 B

A

B

A

B

Rare

Occasional
Common
Abundant

94

75

25

1

24.1

19.5

6.5

0.3

79

75

20

28.0 54

26.6 51

7.1 9

21.2

20.1

3.5

Total

195

50.4

174

61.7 114

44.8

A=Number of records. B=Per cent of total number of
, samples collected at that depth.

47. Ceratium massiliense (Gourret) Jorgensen
Figures 22E-L, chart 38, appendix table 43

Ceratium massiliense is one of the most variable
and difficult species of the genus. The typical form is
shown in figure 22F (which, incidently, is a two -celled
chain). It has the most common form of body, although
the horns frequently are much shorter. Variety armatum
(Karsten) Jorgensen is shown in figure 22L. This was
rare in the Carnegie collection. Peters (1934) consid-
ered it to be a cold-water subspecies in the South Atlantic,
but in our collection no correlation with lower tempera-
ture was evident. Variety protuberans (Karsten) Jorgen-
sen is another variation of the species which has been
recorded in the literature. It has a somewhat elongated
body and thin antaplcal horns, which are somewhat more
widely spread than in the typical form (fig. 221). There
is a form in the Atlantic which simulates somewhat C.
deflexum (fig. 22H). This probably accounts for Peters'
(1934) including C. deflexvun in the species C . massiliense
(see next species below). It will be noted that in this va-
riety of C. massiliense the left antapical horn, where it
leaves the body, is directed slightly antapically so the
posterior curvature of the horn and the posterior edge of
the body are not in the same line. The right antapical is
directed somewhat more posteriorly than in the main
form of the species. This type of divergence, as regards
antapical horns, may be accentuated even more, so that
the specimens resemble, to a degree, C. macroceros (fig.
22G). In this case the curvatures of the antapical horns
extend to a distance about 0.5 transdlameter beyond the
body. A variation of this is shown in figure 22J in which
the right horn does not extend far posteriorly but later-
aUy. Still another variation was found in which the ori-
gins of the antapicals are more lateral than in the main
species, and the horns themselves are much more wide-
spread (fig. 22K).

Most of the Carnegie material of this species could
be placed in one of the above nine varieties. Inter grades
were fairly common, however, so the complex as a whole
represents a single variable species. The varieties
showed no correlation with geographic position or hydro-
graphic conditions, except that the defleximi type was
confined to the Atlantic.

There is also a form which tends toward the body
shape of C. contrarlum (fig. 22E). The left antapical
leaves the body in line with the posterior edge of the body;
the right extends somewhat posteriorly. The horns are
slender. A four-celled chain was found in this form.

The species, as a whole, is one of the most common

CERATIUM SPECIES OF CARNEGIE COLLECTION

39

of the genus. It is a very tolerant tropical species which
may be displaced by currents. It was found at a total of
152 stations--28 in the Atlantic and 124 in the Pacific.
In the Atlantic it was found at practically every station
in the warm Atlantic region and at three stations in the
cold Atlantic region, the most northern of these being
station 6c off Ireland where the surface temperature was
12.°4 C. In the Pacific it occurred almost continuously
throughout the warm Pacific and southeast Pacific re-
gions. It extended northward at either side of the Pacif-
ic, somewhat beyond the limits of this region: off Japan
to station 116 where the surface temperature was 16.°1
C; and off the United States to station 128, surface tem-
perature 16.°4 C. In the southeastern Pacific itoccurred
at the southernmost station, station 60, where the sur-
face temperature was 15.°0 C.

The environmental conditions in situ were: temper-
ature, 10.°6 to 29°5 C; salinity, 29.7 to 37.0 per mille;
pH, 7.17 to 8.47; phosphate, 2 to 233 mg P04/m3.

Of the 476 records, 194 were from water containing
less than 10 mg P04/m3.

The species is quite common, as shown by the num-
ber of sample records which totaled 479. Of these, 159
were rare, 210 occasional, 102 common, and 8 abundant.
Net samples predominated, with 348 records as against
131 pump records. The large number of net records
doubtless was owing to the large spread of the horns
which made the capture of this species efficient in the
coarser cloth of the tow net.

Nielsen (1934) classified C. massiliense as a sur-
face species. The Carnegie data corroborate this (table
40). The species was found with decreasing frequency
with increase in depth although the decrease was not great.

Table 40.

Records of occurrence of C. massiliense
at three levels

Occur -

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional
Common
Abundant

Total

65

96

55

5

16.9

24.9

14.3

1.3

45

65

31

2

15.6

22.3

11.0

0.7

49

49

16

1

18.9

19.3

6.3

0.4

221

57.4 143 49.6 115 44.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

48. Ceratlum deflexum (Kofoid) Jorgensen
Figures 22C-D, chart 39, appendix table 44

Ceratlum deflexum is an intolerant tropical species,
probably restricted to the Pacific and Indian oceans.
Peters (1934), in his study of the material of the Meteor
from the South Atlantic, included C. deflexum as a vari-
ation of C. massiliense. Unfortunately, he did not give a
figure of it. Undoubtedly it was the variety mentioned
above (p. 38, fig. 22H), which is a variant of C. massiliense
but is not to be identified with C. deflexum. Ceratlum
deflexum is characterized by a ventral curvature of the
antapical horns (fig. 22D) which is not found in any of the
variants of C. massiliense . In addition to this, the pos-
terior extension of the horns is much greater than in C^
massiliense. Ceratlum deflexum is a distinctive species
and is easily recognized after it has been observed once.

In the Carnegie collection it was found at fifty-three
stations--all in the Pacific. There were 105 records of
occurrence, of which 58 were rare, 34 occasional, and
13 common. Eighty-four were from net records and 21
were from pump records.

Most of the stations were concentrated in the east-
ern and central Pacific between latitudes 20° north and
20° south, except for one station at Japan and two sta-
tions in the southern part of the southeast Pacific region.
The species probably is not as limited in its distribution
as the Carnegie data indicate, since it was found off San
Diego, California by Kofoid (1907a), in the China Sea
south of 20° north latitude by Bohm (1931), and in the
Indian Ocean by Jorgensen (1911).

The species was found only at stations with high
temperatures, except at the two stations to the southeast.
The surface temperatures at the main group of stations
ranged from 18.°7 to 29.°3 C; at the two stations in the
southeast, 16.°9 and 19.°2 C. The ranges of environmen-
tal conditions in situ were: temperature, 10.°8 to 29.°3C;
salinity, 29.7 to 36.5 per mille; pH, 7.82 to 8.47; phos-
phate, 4 to 166 mg P04/m3.

The curious distribution of this species may be ow-
ing to a preference for eutrophic water. In the large
oligotrophlc area of the North Pacific, where the average
phosphate content of the upper 50 meters was less than
10 mg P04/m3, there were only nine stations. Its ab-
sence off Peru might be attributed, then, to lower tem-
peratures, and its absence in the Easter Island region to
oligotrophlc conditions. The greatest number of stations
were in the regions where the average phosphate content
was above 25 mg and the greatest populations were found
at stations where the phosphate content was above 25 or
50 mg P04/m3. In this connection it is to be noted that
of the 107 sample records for the species, only 11 were
in water containing less than 10 mg P04/m3.

The species is a surface form. In the Carnegie col-
lection it was found with decreasing frequency with in-
crease in depth (see table 41). Thus, it was found in 15.5
per cent of the surface samples and only 6.7 per cent of
the 100-meter samples. Because of contamination of the
deeper hauls, the real distribution is probably even more
definitely a surface one than the present data indicate.

Table 41.

Records of occurrence of C. deflexum
at three levels

Occur-
rence

Depth in meters

50

100

A B

A

B

A B

Rare

Occasional

Common

32

19
11

8.0
4.7
2.8

16

11

1

5.7
3.9
0.3

11
5

1

4.3
1.9
0.4

Total

62

15.5

28

9.9

17

6.6

A=Number of records. B=Per cent of total number of
samples collected at that depth.

49. Ceratlum carriense Gourret
Figure 22A, chart 40, appendix table 45

This is a rather variable species of uncertain taxo-
nomic position. Certain varieties have been named which
show characteristic spread of horns (for instance, var.
volans Cleve and f. cevlonicum SchrSder), but in the

40

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Carnegie collection many intergradations were found
between these and still other variants so they have not
been recorded. Some of the variants are similar to
modifications of C. massiliense. Indeed, these two spe-
cies may be identical. They have been kept separate in
this report on the basis of the origin of the left antapical
horn which in C. carriense starts in a posterior direc-
tion before curving laterally, so at its proximal end it
does not form a straight line with the posterior edge of
the body. The demarcation was indefinite, however, with
numerous inter grades.

The distribution of the species as considered here Is
similar to that of C. massiliense . except that it does not
extend as far outside the tropical regions. In the Carne-
gie collection in the Atlantic there is only one station
outside the warm Atlantic region, station 13a. In the
Pacific it has about the same distribution as C. massili-
ense in the north, but was not well represented in the
southeastern part of the Pacific (chart 40). Nielsen
(1934) found it in the Pacific from Panama to Australia,
and Peters (1934) found it widely distributed in the warm
water of the South Atlantic.

The total number of Carnegie stations at which it was
found, was 111, 12 of which were in the Atlantic and 99 in
the Pacific. The total number of sample records for the
species was 281, with 138 rare, 124 occasional, and 19
common. The net records numbered 219; the pump rec-
ords, 62.

The range of environmental conditions in which the
species was found was similar to that of C. massiliense .
though somewhat more restricted. The surface temper-
atures at its record stations in the Atlantic ranged from
21.°2 to 28.°5 C and in the Pacific from 16.°3 to 29.°5 C.
Of the 281 records, 98 were from water containing less
than 10 mg P04/m3. The complete ranges of environ-
mental conditions in situ were: temperature, 12.°5 to
29.°4 C; salinity, 29.7 to 35.9 per mille; pH, 7.82 to 8.47;
phosphate, 3 to 153 mg P04/m3.

Ceratixim carriense is a surface species (Nielsen,
1934). In the Carnegie collection It was found with de-
creasing frequency with increase in depth (table 42).

Table 42.

Records of occurrence of C. carriense
at three levels

Occur-
rence

Rare

Occasional

Common

Total

Depth in meters

B

50

B

100

58
51
13

13.5

13.0

3.4

47
43

14.4

14.2

2.2

33
30

12.0
11.9

122 29.9 96 30.8 63 23.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

50. Ceratium contrarlum (Gourret) Pavlllard
Figures 22E, 24A-B, chart 41, appendix table 46

This species Is rather distinctive, and fairly con-
stant in its body shape and origin of horns. There is one
variation of the species, however, which has club-shaped
ends on the antapical horns, var. claviceps Schroder (fig.
24B). In this variant the apical horn also maybe expand-
ed, but only gradually toward the tip. It was found rather

frequently in both the Atlantic and Pacific. It apparently
does not represent a response to shade conditions, as it
was found frequently at the surface.

The species as a whole is one of the most common
of the genus. Peters (1934) found it commonly in the
South Atlantic to about 33° south latitude generally and
to almost 40° south, south of the Cape of Good Hope.
Nielsen found it at all oceanic stations in the South Pa-
cific with surface temperatures over 19° C. In the
Carnegie collection it occurred at almost all the stations
in the warm Atlantic region, warm Pacific, and southeast
Pacific region as well as at one station in the cold North
Pacific region (chart 41). The total number of station
records was 138, of which 20 were in the Atlantic and
118 in the Pacific. The total nimiber of sample records
was 467, of which 167 were rare, 172 occasional, and
128 common. The net records numbered 298; the pump
records, 169.

The distribution of the species at Carnegie stations
indicated a greater tolerance for lower temperatures in
the Pacific than in the Atlantic. In the Atlantic it did not
occur in the North Atlantic Drift but stopped abruptly at
the 20° isotherm. In the Pacific, on the other hand, it
occurred to surface temperatures of 16.°4 off the United
States, and 15° C at 40° south in the southeastern Pacific.
It showed no regard for the nutrient content of the water,
being foimd as much in oligotrophic as eutrophic water.
Of the 467 records of occurrence, 154 were in water con-
taining less than 10 mg P04/m3. The environmental
conditions in situ were as follows: temperature, 10.°2
to 29.°5 C; salinity, 30.7 to 37.1 per mille; pH, 7.76 to
8.47; phosphate, 2 to 178 mg P04/m3.

Ceratium contrarlum is a surface species. Nielsen
(1934) always found it more common in the 50- to 0-meter
samples than in the deeper samples. In the Carnegie col-
lection it was found with decreasing frequency with in-
crease in depth (table 43). For example, it was found In
57.2 per cent of the surface samples, but only 37.4 per
cent of the 100-meter samples.

Table 43.

Records of occurrence of C. contrarlum
at three levels

Occur-
rence

Depth in meters

50

100

A B

A B

A B

Rare

Occasional

Common

80
83
58

20.8
21.5
15.1

50
60
41

17.4
21.3
14.5

37
29
29

14.6
11.4
11.4

Total

221

57.4

151

53.2

95

37.4

A=Number of records. B=Per cent of total number of
samples collected at that depth.

51. Ceratium trichoceros (Ehrenberg) Kofoid
Figure 22B, chart 42, appendix table 47

This Is a widespread intolerant tropical species
characterized by widespread horns which lie parallel in
their distal parts. It is fairly constant in its shape, al-
though Nielsen (1934) has noted a tendency for an increase
in diameter in colder or neritic waters.

In the Carnegie collection the species was found wide-
ly distributed in the warm regions, although not so com-
monly as such species as C. massiliense. Ceratlxim

CERATIUM SPECIES OF CARNEGIE COLLECTION

41

trichoceros occiirred at eighty-six stations- -fourteen in
the Atlantic and seventy-two in the Pacific. The total
number of sample records for the species was only 181,
of which 89 were rare, 67 occasional, 24 common, and 1
abundant. The net records numbered 118; the pump rec-
ords, 63. Peters (1934) found it widespread in the warm
South Atlantic but principally In his region I and along
the west coast of Africa. Nielsen (1934) found it through-
out the region from Panama to Australia.

According to the Carnegie distribution this is a
rather strictly tropical species. In the Atlantic it was
confined to the warm Atlantic region, with surface tem-
peratures from 21.°2to 28.°5 C. In the Pacific it was con-
fined to the warm Pacific region and the southeast Pa-
cific region. The total range of surface temperatures in
the Pacific was from 19.°0 to 29.°5 C. The total ranges
of environmental conditions in situ were: temperature,
14.°0 to 29.''4 C; salinity, 31.7 to 37.1 per mille; pH, 7.82
to 8.47; phosphate, 2 to 159 mg P04/m3.

The Carnegie data do not show any preference on
the part of this species for any particular type of water
within the warm-water regions. It does not shun oligo-
trophlc water. Of the 181 records of occurrence, 63
were from water containing less than 10 mg P04/m3.

It is uncertain at what depth this species normally
lives most abundantly. Nielsen (1934) stated that it was
always more abundant in his 50- to 0-meter samples. In
the Carnegie collection, however, there were more rec-
ords for 100 meters than for the upper levels, with the
fewest records for the svtrface (see table 44).

Table 44.

Records of occurrence of C. trichoceros
at three levels

Occur-
rence

Depth in meters

50

100

A 1 B

A 1 B

A 1 B

Rare

28

7.3

23

8.1

38

15.0

Occasional

20

5.2

23

8.1

24

9.5

Common

8

2.1

10

3.6

6

2.4

Abundant

1

0.3

Total

57

14.9

56

19.8

68

26.9

A=Number of records. B=Per cent of total nimiber of
samples collected at that depth.

52. Ceratium vultur Cleve
Figures 23A-H, chart 43, appendix table 48

The species as here considered Includes C. simiatra-
num (Karsten) Jorgensen and C. pavlllardii Jorgensen.
Ceratium vultur is a variable species with many variable
characters giving rise to what seem, at times, separate
species. The study of the extensive material of the Car-
negie collections, however, convinced the authors that
these are variants of a single large species. The inter-
grades between them are numerous. Peters (1934) and
Nielsen (1934) have already considered C. vultur and C,
sumatranum as a single species, with sumatranum a va-
riety of the former. In our material it was possible to
distinguish seven different varieties as follows.

Variety vultur (Cleve) is stout and long-horned (fig.
23H). The right antapical ascends moderately; the left
antapical extends posteriorly less than 0.5 girdle width
before turning anteriorly.

Variety iaponicum (Schroder) is as above, except that

the left antapical extends posteriorly more than 0.5 girdle
width before turning anteriorly (figs. 23E-F).

Variety sumatranum (Karsten). The right antapical
extends more or less laterally. The left antapical ex-
tends posteriorly a varying distance before turning an-
teriorly (fig. 23D).

Variety pavlllardii (Jorgensen). The right antapical
is as in var. sumatranum . The left antapical is kinked at
the origin, and turns abruptly to the anterior (fig. 23C).

Variety regulare n. var. The right antapical extends
laterally at the base but turns anteriorly. The left ant-
apical does not extend posteriorly, but laterally, before
turning anteriorly. In this respect it resembles C.
masslllense (fig. 23G).

Variety reversum n. var. The right antapical is
kinked at the origin and turns abruptly to the anterior.
The left antapical extends somewhat posteriorly before
turning anteriorly. The curvature of the antapical horns,
then, is the reverse of that in var. pavlllardii (fig. 23B).

Variety recur vum Jorgensen. The right antapical
begins laterally, then at about half its length turns pos-
teriorly. The left antapical begins posteriorly, turns
laterally, and then posteriorly (fig. 23A).

Most of these varieties had the same distribution in
both the tropical Atlantic and Pacific oceans. Varieties
regulare and reversum . however, were confined to the
Pacific, whereas var. recur vum was common in the North
Equatorial Current of the Atlantic but occurred at only
two stations In the entire Pacific, stations 98 and 149.

The species as a whole is intolerant tropical and
confined to regions of warm water. Peters (1934) found
C. vultur to be confined to warm, definitely oligotrophic
water. Nielsen (1934) found it in the Panama region and
the South Equatorial Current. In the Carnegie collection
in the Atlantic it was confined to the warm Atlantic re-
gion, with surface temperatures ranging from 20.°5 to
28.°5 C. It was found at all stations in this region except
one (chart 43). In the Pacific it was confined to the warm
Pacific and southeast Pacific regions. The surface tem-
peratures at the Pacific stations varied from 19.°4 (sta-
tion 66 in the southeast Pacific) to 29. °4 C. The ranges
of environmental conditions in situ were: temperature,
19.°8 to 29.°3 C; salinity, 29.7 to 37.1 per mille; pH, 7.76
to 8.42; phosphate, 2 to 189 mg P04/m3.

It occurred at ninety-nine stations- -twenty-four in
the Atlantic and seventy-five in the Pacific. There were
251 records of occurrence, with 168 rare, 71 occasional,
and 12 common. Of these, 203 were net records and 48
were pump records.

Nielsen (1934) foimd both the main species and var.
sumatranum to be shade species. The Carnegie data
substantiate these findings in that they show an increasing
frequency with Increase in depth (table 45).

Table 45

Records of occurrence of C.
at three levels

vultur

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

B

Rare

Occasional

Common

58

27
8

15.1
7.1
2.1

59
21

1

20.1
7.5
0.4

51

23

3

20.1
9.1
1.2

Total

93

24.3

81

28.0

77

30.4

A=Number of records. B=Per cent of total number of
samples collected at that depth.

42

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

53. Ceratium horridum Gran

Figures 23I-L, 24C-I, 25A-G,

chart 44, appendix table 49

This species comprises a variable and uncertain
group of the genus. The authors cannot agree with Jor-
gensen (1920), however, who combines this group withC^
tenue . Peters (1934) followed Jorgensen's classification
but did not name or figure any of the forms of the inclu-
sive species. Paulsen (1930) and Nielsen (1934), on the
other hand, separated C. horridum from C. tenue . as the
authors have done in the present report. The two spe-
cies can be separated on the basis of body shape and cur-
vature of horns and also, in part, on the basis of size.
The different shapes can be seen by reference to figures
23 to 26. Ceratium horridum is larger than C. tenue .
having an average diameter of 44 microns as compared
with 37 microns for C. tenue. The extremes, however,
overlap, with a minimum of 30 microns for C. horridum
and a maximum of 45 microns for C. tenue.

Ceratium horridum is made up of a number of vari-
able units; formerly separated as C. horridimi (C. inter -
medium) . C. moUe . and C. claviger . Nielsen (1934) still
considered C. claviger a species distinct from C. molle .
but in the Carnegie collection no evidence for this sepa-
ration was found. It seems best, therefore, to treat
these forms as varieties.

Variety molle and var. claviger are tropical forms,
whereas var. horridum is more or less cosmopolitan. It
is probable that there are northern and tropical forms of
var. horridum which overlap in such a way that they are
not distinguishable. In any event, a certain difference
can be discerned between the cold- and warm -water
forms. In figures 24D and G are two specimens repre-
sentative of the cold-water form of the North Atlantic;
figure 24H shows a specimen from the warm Gulf Stream
or North Atlantic West Wind Drift. Peters (1934) found
the inclusive species widespread over the South Atlantic,
as did Nielsen (1934) in the South Pacific.

The Pacific material is similar to that of the Atlan-
tic, at least in part. The variety in the Panama region,
for instance, (fig. 24E) is similar to the Gulf Stream
form. The Galapagos form (fig. 25B), on the other hand,
rather closely resembles the cold-water form of the
North Atlantic. In the cool water of the southeast Pacific
the variety was more robust and long-horned (fig. 25E).
In the Humboldt Current the variety was even more ro-
bust and the horns were more extended (fig. 25F). In the
warm equatorial waters it was slender and long-horned
(fig. 24C). In the cold waters of the cold North Pacific
region it was found only at three stations off the west
coast of North America, stations 126 to 128, and at one
station off Japan, station 114 (figs. 241, 25A). At these
stations the variety did not resemble the cold-water form
of the North Atlantic. It was slender and long-horned,
resembling much more the equatorial form of the Pacific
(figs. 24C, 25G). It was, however, much more spiny.
The surface temperatures at the eastern stations varied
from ll.°2 to 16.°4 C and at the station off Japan It was
19.°9 C. The differences between the forms of var. hor-
ridum in the northern Atlantic and northern Pacific is
another indication of the Isolation of the northern Pacif-
ic waters. Its absence in the most northern (coldest)
stations in this region, however, may be owing to the
temperat\ires, which were mostly less than 10° C. This
was lower than the temperatures of the northern Atlan-
tic where the variety occurred.

Variety molle is simil<i.- to var. horridum but is
confined to the warm regions. Its occurrence in the Car-
negie collection was sporadic. Its most northern position
in the Atlantic was at station 3 (see fig. 25D). Here it
was rather robust but had the typical long horns. In the
warmer stations it was more slsnder and delicate (see
fig. 231). The Pacific form of var. molle was quite sim-
ilar to that of the Atlantic (see fig. 23K).

Frequently varieties were found which were difficult
for the authors to assign to either molle or horridum.
Figures 25C and G show two such. In a sense these are
intergrades between the two varieties, but figure 25G
shows a specimen with extremely long and attenutated
antapical horns. Specimens such as this have been part-
ly responsible, perhaps, for the opinion that C. horridum
and C. tenue are part of the same species complex. It is
not likely, however, that this represents an intergrade,
as the origin of the horns and proximal curvatures are
different from those of C. tenue (cf. figs. 26C-D).

Among recent workers, Melsen (1934) is the only
one who considered C. claviger Kofoid a separate spe-
cies. This opinion probably is owing to a lack of suffi-
cient material, as it is plain from the Carnegie material
that this form Is characterized only by the swollen ends
of the antapical horns. The body shape and origins of
horns may be that of var. horridum or var. molle (see
figs. 23J, L),

The species as a whole Is not very common. It was
found at a total of eighty Carnegie stations- -twenty-eight
in the Atlantic and fifty-two in the Pacific. Since there
are apparently cold- and warm -water varieties of the
species, the ranges of temperatures for these stations
are not very significant. In the Atlantic the siu^ace tem-
peratures at the record stations varied from 8.°9 to 28.°5
C and in the Pacific from ll.°2 to 29.°4 C. The environ-
mental conditions In situ were as follows: temperature,
6.°3 to 29.°3 C; salinity, 31.7 to 36.8 per mille; pH, 7.68
to 8.39; phosphate, 4 to 233 mg P04/m3.

The total number of sample records for the species
was only 149, of which 96 were rare, 38 occaslona4, 11
common, and 4 abimdant. The net records numbered 96;
the pump records, 53.

Ceratium horrid\mi is not a species of oligotrophlc
water. Although it was found in some "barren" regions,
It occurred predominantly in richer waters. Only 16 of
the 149 records were in water containing less than 10 mg
P04/m3.

The species is a decided shade form. In the cold-
water region of the North Atlantic it was found about
equally at the three levels, but elsewhere it was found
much more frequently at greater depths, particularly at
100 meters. The depth records for the species throughout

Table 46. Records of occurrence of C
at three levels

. horridum

Occur-
rence

Depth in meters

50

100

A

B

A

B

A

1 B

Rare

Occasional
Common
Abundant

31
12

1
3

8.1
3.1
0.3
0.8

32

14
3
1

11.4
4.9
1.1
0.3

33

12

7

13.0
4.7
2.8

Total

47

12.3

50

17.7

52

20.5

A=Number of records. B=Per cent of total number of
samples collected at that depth.

CERATIUM SPECIES OF CARNEGIE COLLECTION

43

its entire range show an increase in frequency with an
increase in depth (table 46).

54. Ceratium tenue Ostenfeld and Schmidt
Figures 26C-D, chart 45, appendix table 50

Two varieties of this species can be recognized:
var. Inclinatum . in which the antapical horns are direct-
ed more or less apically (fig. 26C); and var. tenuissi-
mum . in which the antapical horns are directed laterally
(fig. 26D). It is important to note that the origin and
proximal curvatures of the antapicals in these varieties
are the same, the differences being due to ciu"vatures
more than one-half girdle diameter from the body.

The two varieties do not have separate ranges of
distribution but are found together in the warm regions
of the world. The species as a whole is more common
than C. horridura . but does not have any cold-water rep-
resentative.

Peters (1934) apparently included this form in C.
horridum . following Jorgensen (1920), but he gave no fig-
ures. Nielsen (1934), who separated it from C. horridxun .
stated that it had a distribution in the South Pacific sim-
ilar to that species.

The total number of station records for the species
in the Carneple collection was 100--19 of these were in
the Atlantic and 81 were in the Pacific. The Atlantic
records were all restricted to the warm Atlantic region,
with surface temperatures from 21. °2 to 28.°5 C. The
Pacific stations were in the warm Pacific region and the
southeast Pacific region, except for one station off Cali-
fornia which extended Into the cold Pacific region (sta-
tion 128, surface temperature 16.°4 C). Throughout its
range the species was found fairly uniformly except off
Peru and the Panama and Galapagos areas, where it sel-
dom occurred.

The total number of sample records for the species
was 236, of which 129 were rare, 99 occasional, and 8
common. The net records numbered 115; the pump rec-
ords, 121. The environmental conditions in situ were as
follows: temperature, 8°8 to (29.°4) C; salinity, 33.2 to
37.1 per mllle; pH, 7.93 to 8.39; phosphate, 2 to 176 mg
P04/m3.

Ceratium tenue Is definitely a shade species. In the
Carnegie collection it was foimd with increasing frequen-
cy with increase in depth. There were almost four times
as many records for 100 meters as for the surface (table
47). Nielsen (1934) also foimd the species to be a shade
form in the Pacific.

Table 47.

Records of occurrence of C. tenue
at three levels

Occur-
rence

Depth in meters

50

100

A B

A B

A B

Rare

Occasional

Common

29

14

1

7.3
3.6
0.3

43
35

1

14.2

12.1

0.4

57

50

6

20.9

18.9

2.4

Total

44

11.2

79

26.7

113

42.2

A=Number of records. B=Per cent of total nimiber of
samples collected at that depth.

55. Ceratium longissimum (Schroder) Kofoid
Figures 26A-B, chart 46, appendix table 51

This is a rare intolerant tropical species. It is not
variable in form and for this reason it is interesting to
note a constant difference in the spread of the horns be-
tween the Atlantic and Pacific specimens. In the Atlantic
form (fig. 26A) the three horns are practically parallel,
whereas in the Pacific form (fig. 26B) the antapicals, al-
though proximally almost parallel in position, soon di-
verge considerably from the apical horn. Also, the horns
In the Pacific form are much more slender.

The species was found at twenty-eight stations- -five
in the Atlantic and twenty-three in the Pacific. The At-
lantic stations were all In the warm Atlantic region, with
surface temperatures above 20° C. The Pacific stations
were in the warm Pacific and southeast Pacific regions
with siu-face temperatures above 21° C. There was a
particular concentration of stations in the general area
between Hawaii and North America. The distribution as
a whole was closely restricted as to latitude, lying be-
tween latitudes 14° south and 34° north. Nielsen's (1934)
records, which were the first for the Pacific, lie within
these limits except for one station In Australian waters
at latitude 32° south. The temperature there, however,
was above 20° C. In the Atlantic the northern limit was
somewhat farther north than in the Pacific, namely, 38.°5
north.

The rarity of the species is further shown by the
nimiber of sample records, which was thirty-two. Of
these, twenty-five were rare and seven occasional. Only
six were pimip records; twenty-six were net records.
The species may prefer oligotrophlc water as it was sel-
dom found in regions rich in plankton and fifteen of the
thirty-two records of occurrence were in water contain-
ing less than 10 mg P04/m3. The ranges of environmen-
tal conditions in situ were: temperature, 12.°3 to 28f0 C;
salinity, 34.3 to 36.4 per mllle; pH, 7.92 to 8.39; phos-
phate, 3 to 138 mg P04/m3.

Ceratium longissimum is one of the most pronounced
shade species encountered. Of the thirty-two records of
occurrence, none were for the surface, only five for 50
meters, and twenty-seven for 100 meters 'see table 48).
Nielsen (1934) found it only in the samples taken from 200
to 100 meters and 100 to 50 meters.

Table 48. Records of occurrence of C. longissimum
at three levels

Occur-

Depth

in meters

50

100

A

B

A

B

A 1 B

Rare
Occasional

5

1.8 20 7.8
7 2.8

Total

5

1.8 27 10.6

A=Number of records. B=Per cent of total number of
samples collected at that depth.

56. Ceratium arctlcum (Ehrenberg) Cleve
Figures 26E-H, 27A-E, chart 47, appendix table 52

Ceratium longloes (Bailey) Gran and C. arctlcum
(Ehr.) Cleve Intergrade in such a way that the authors are
forced to include them under a single species. According

44

CERATIUM m THE PACIFIC AND NORTH ATLANTIC OCEANS

to the international rules of botanical nomenclature the
name for this Inclusive species must be C. arctlcum. as
it was used as a specific name earlier than C. longlpes
(see Jorgensen, 1911, pp. 84-85). The longlpes form,
however, is sufficiently distinct to be separated as ava-
riety. Variety ventricostmi Ostenf eld is a robust, heav-
Uy armored form, with swollen body (fig. 26E). It is
apparently limited to the northern Atlantic waters. In
the Carnegie collection it occurred only in the cold At-
lantic region, with the single exception of station 14,
where the surface temperature was 21.°2. At the other
record stations the temperatures were under 11° C.

Variety arcticimi probably is more characteristic of
colder water than is longlpes . although not so much so
as the name would indicate. Variety longlpes is found
more frequently over a wider range than var. arcticum .
but the latter strays far from its northern home. Cleve
(1900) found it as far south as the Azores and the Straits
of Glbralter. In the Carnegie collection it was found as
far south as 20° north in the central Atlantic (station 20)
and at one station in the Caribbean (station 33). It was
never found at the surface in these warm regions, how-
ever, although the temperatures In situ were quite high;
namely, 25.°8 C at station 20 and 28.°2 C at station 33.
This is a curious distribution for a species of Ceratiimi.
Its center of abundance unquestionably is in the cold
waters where the temperatures are less than 10° C at
the surface in the summer; yet isolated specimens find
their way into the heart of the tropics.

Apparently a similar situation exists in the North
Pacific. The main distribution of arcticum is to the
north in cold water ( Carnegie stations 120 to 121, sur-
face temperatures less than 8° C), yet a specimen was
foimd at station 146 between Hawaii and the United States,
where the surface temperature was 22.°4 C. To be sure,
the record was for the 100-meter sample, but even here
the temperature was 19.°7 C.

Variety longlpes . on the other hand, although it oc-
curs abundantly over a wider range than var. arcticum .
is not found so far from its centers of abundance as var.
arcticum.

The inclusive species was foiind at a total of thirty-
one stations, of which twenty-one were in the Atlantic
and ten in the Pacific. The total number of sample rec-
ords was eighty-four, with thirty-seven rare, twenty-
seven occasional, eleven common, and nine abundant.
Net samples predominated with fifty-nine records, as
compared with twenty-five pump sample records.

The total ranges of environmental conditions in situ
V'ere: temperature, -l.°2 to 28.°2 C; salinity, 32.7 to
37.1 per mille; pH, 7.64 to 8.29; phosphate, 3 to 228 mg
P04/m3.

No one has reported anything regarding the depth at
which the species normally lives since Nielsen (1934) did
not find the species. The Carnegie records (table 49)
show an Increase in frequency with increase in depth to
100 meters. The increase is not great, however, with
the species occurring in 8.1 per cent of the surface sam-
ples and 10.6 per cent of the 100-meter samples. Is it
possible that C. arcticum . which is characteristic of arc-
tic waters, is a shade species adapted for life under the
polar Ice cap where the intensity of illumination cannot
be great? The Carnegie data alone cannot be conclusive
regarding the vertical distribution of this species. Many
of the station records are from cold northern waters
where the plankton In the upper layers is comparatively
dense so there is much greater chance of contamination

of the open nets dxiring hauling in. On the other hand. It
should be noted that the records for the species in the
warm oligotrophlc water of the Atlantic are predominant-
ly subsurface records. This occurrence may be owing to
the lower temperatures obtaining there rather than to the
preference for, or superior survival in, levels of weak
illumination so the species may not be a shade species
In the true sense of the term.

Table 49.

Records of occurrence of C. arcticum
at three levels

Occur -

Depth in meters

50

100

A

B

A

B

A

B

Rare

7

1.8

14

5.0

16

6.3

Occasional

12

3.2

7

2.5

8

3.2

Common

6

1.6

4

1.4

1

0.4

Abundant

6

1.6

1

0.4

2

0.8

Total

31

8.2

26

9.3

27

10.7

A=Number of records. B=Per cent of total number of
samples collected at that depth.

57. Ceratium hexacanthum Gourret
Figures 27F-G, chart 48, appendix table 53

This is a common, very tolerant tropical species.
The antapical horns tend to be extremely thin and atten-
uated, and to curl so that various modifications of these
horns occur. There seems to be no justification for cre-
ating varieties such as var. spiralis Kofoid, on the basis
of such characters as the curl of the horns.

In the Pacific the species was confined to the warm
tropical waters. The surface temperatures at the Pacif-
ic record stations were all above 20° except at two sta-
tions in the southeastern Pacific (stations 57 and 62),
where the surface temperatures were 19.°0 and 19.°2 C.
In the Atlantic, on the other hand, the species is appar-
ently carried by the North Atlantic Drift to the British
Isles and as far as Iceland (see chart 48) Into water with
surface temperatiu-es as low as 8.°9 C. Morpholgically
no separation could be made between the Atlantic and
Pacific forms. The form carried into the cold waters of
the North Atlantic, however, probably is a particular sub-
species, at present indistinguishable from the tropical
form because the tropical form does not invade cold
water elsewhere. Peters (1934) stated that it was no-
ticeably absent from the South Atlantic south of about 33°
south. It dropped out at about this latitude at the Carne -
gie stations in the southeastern Pacific, and at the Dana
stations in the southeastern Pacific (Nielsen, 1934).
Thus, it kept within the 20° isotherm in the North Pacif-
ic, which indicates that the race of the Pacific is at least
physiologically different from that in the North Atlantic.

The ranges of environmental conditions in situ for
the species as a whole were: temperatiure, 8.°2 to 29.°3
C; salinity, 30.7 to 37.1 per mille; pH, 7.17 to 8.47;
phosphate, 2 to 198 mg P04/m3.

The total number of station records was 124- -28 in
the Atlantic and 96 in the Pacific. There were 327 rec-
ords of occurrence, of which 169 were rare, 129 occa-
sional, and 29 common. The net records numbered 267;
the pump records, 60.

Nielsen (1934) classified C. hexacanthum as a shade

CERATIUM SPECIES OF CARNEGIE COLLECTION

45

species. He found tiiat in the eastern part of the South
Pacific, where there were stations rich in plankton, the
species had its main distribution above 50 meters; at
the western stations, where the plankton population was
less dense, the species had its main distribution below
50 meters. The Carnegie data do not, however, substan-
tiate these results. The number of records in the Car-
negie collection decreased with increase in depth. Per-
haps, if a classification of the Carnegie tropical stations
as to plankton content could be made, some correlation
with plankton density could be discovered. The authors
can conclude that C. hexacanthum . however, is not a
typical shade species as many other species in the Car-
negie collection showed a decided increase in frequency
of occurrence with increase in depth, considering the
collection as a whole.

Table 50.

Records of occurrence of C.
at three levels

hexacanthum

Depth in meters

rence

50

100

A 1 B

A B

A B

Rare

Occasional

Common

Total

72
63
13

18.2

16.3

3.4

53

43

9

17.9

15.2

3.2

44 17.3

?3 9.1

7 2.7

148

37.9 105 36.3 74

29.1

A=Number of records. B=Per cent of total number of
samples collected at that depth.

58. Ceratlum reflexum Cleve
Figure 27, chart 49, appendix table 54

This is a very rare, intolerant tropical species. It
was not reported from the South Atlantic by Peters (1934)
although it had been reported from as far south as 19.°5

south latitude in the Atlantic by Cleve (1900). Nielsen
(1934) found it at only four stations in the South Pacific.
In the Carnegie collection it occurred at twenty-one
stations--six in the Atlantic and fifteen in the Pacific.
The Atlantic stations were between 11° and 24° north
latitude. The Pacific stations were much scattered (see
chart 49). The range of surface temperatures at the rec-
ord stations for the species was from 23.°4 to 29.°4 C.
The ranges of environmental conditions in situ were:
temperature, 17.°5 to 28.°6 C; salinity, 34.4 to 37.1 per
mille; pH, 7.99 to 8.39; phosphate, 3 to 123mgP04/m3.

The total number of sample records was thirty-nine,
of which thirty -three were rare, five occasional, and one
common. The record of common was at station 105 near
Guam; the occasional records were from station 20 (trop-
ical Atlantic), station 97 (near the Phoenix Islands), and
station 107 (near Guam). The species perhaps prefers
oligotrophic water since twenty -three of the thirty -nine
records were in water containing less than 10mgPO4/m3.
^ It Is probably a shade species. Practically all Niel-
son's (1934) records were from the deeper layers. The
Carnegie data show an increase in frequency with in-
crease in depth (table 51).

Table 51.

Records of occurrence of C. reflexum
at three levels

Occur-
rence

Depth in meters

50

100

A

B

A

B

A 1 B

Rare

Occasional

Common

8
2

P

2.1
0.6

13

1

4.6
0.4

12
3

4.7
1.2

Total

10

2.7

14

5.0

' 15

5.9

A=Number of records. B=Per cent of total number of
samples collected at that depth.

LITERATURE CITED

Bohm, A. 1931. Distribution and variability of Ceratl-
um in the northern and western Pacific. Bishop
Mus. Bull., no. 87, pp. 1-46, figs. 1-38, pi. 1.

Cleve, P. T. 1900. The seasonal distribution of AUantic
plankton organisms. Goteborgs K. Vetensk. och
Vitterhetssamhailes. Handl., ser.4, vol. 3, 368 pp.

Gourret, P. 1883. Sur les P^ridlniens du golf e de Mar-
seille. Ann. du Mus^e d'hist. nat. de Marseille.
Zoologle T. 1, pp. 1-114, pis. 1-4.

Gran, H. H. 1912. to J.Murray and J.Hjort. The depths
of the ocean, pp. 307-386, figs. 212-252.

Hardy, A. C. 1935. The plankton of the South Georgia
whaling grovmds and adjacent waters, 1926-1927.
Discovery Reports, vol. 11, pt. 5, pp. 273-370.

JSrgensen, E. 1911. Die Ceratien. Eine durze Mono-
graphie der Gattung Ceratium Schrank. Internal.
Rev. gesamt. Hydrobiol. u. Hydrogr., vol. 4, supp.
124 pp., 10 pis.

1920. Mediterranean Ceratia. Rept. Danish

Oceanogr. Exped. 1908-1910 to the Medlter. and
adjacent seas, vol. 2, jour. 1, pp. 1-110, 94 text
figs., 26 charts in text.

Karsten, G. 1907. Das Ihdische Phytoplankton. Wis-
sensch. Ergebn. Deut. Tiefsee-Exped., vol. 2,
pp. 221-548, pis. 34-54.

Kofold, C. A. 1907a. Dinoflagellataof the SanDiego re-
gion, m. Descriptions of new species. Univ. Calif.
Publ., Zool., vol. 3, pp. 299-340, pis. 22-33.

— ; 1907b. The plates of Ceratium with a note on

the unity of the genus. Zool. Anzeig., vol. 32, pp.
177-183, 8 figs.

Lebour, M. 1925. The dinoflagellates of northern seas.
Plymouth Marine Biol. Lab., 192 pp., 35 pis., 53
text figs.

Nielsen, E. S. 1934. Untersuchengen uber die Verbrei-
tung, Biologie und Variation der Ceratien im Sud-
lichen Stillen Ozean. Dana Report No. 4, pp. 1-67,
74 text figs., 24 tables, 11 charts.

Okamura, K. 1912. Plankton organisms from Bonito

fishing grounds. Rept. Imp. Bur. Fish. Japan, Sci.

Invest., vol. 1, pp. 1-35, pis. 1-5.
Paulsen,©. 1908. Peridiniales. In Brandt, K., and Ap-

stein, C, Nordisches Plankton (Kiel and Leipzig,

Lipsius and Tischer), no. 18, 124pp., 155textfigs.
1930. Etudes sur le mlcroplancton de mer

d'Alboran. Trabajos N. 4 Inst. Espanol Oceanogr.,

108 pp., 61 figs. (1930). Madrid.
Pavillard, J. 1905. Recherches sur la flore pelagique

(phytoplankton) de I'Etang de Thau. Trav. Inst. bot.

Univ. Montpellier et Stat. zool. de Cette, Montpel-

lier, s6r. mlxte mdm.2, 116 pp., 1 chart, 3 pis.
1923. A propos de la syst^matique des P^ri-

diniens (2^ note). Bull. Soc. Bot. France, vol. 70,

pp. 914-918, 3 figs.
Peters, N. 1934. Die Bevolkerung des Sudalantischen

Ozeans mit Ceratien. Wissensch. Ergebn. Deut.

Atlantische Exped. Meteor . 1925-1927, vol. 11, pp.

1-69, 4 pis., 28 charts, 11 tables.
Schott, G. 1935. Geographie des Indischen und Stillen

Ozeans. 413 pp., 37 pis., 114 figs. Hamburg.
Schroder, B. 1911. Adriatische Phytoplankton. SitzBer.

Kaiserl., Akad. Wissensch., Wien, math. -nat. Kl.,

vol. 120, pt. 1, pp. 601-657, 16 text figs.
Shiviroff, P. and E. Fedorov. 1938. Scientific work of

the drifting North Polar Station. Nature, vol. 141,

pp. 629-632.
Sverdrup, H. U., F. M. Soule, J. F. Fleming, and C. C.

Ennis. 1944. Observations and Results in Physical

Oceanography. Oceanogr. I-A and I-B. Carnegie

Inst. Wash. Pub. No. 545, I-A, 156 pp., 48 figs.;

I-B, 315 pp.

46

APPENDIX TABLES 1-54
TABLES OF DISTRIBUTIONAL AND ENVIRONMENTAL RECORDS

Abbreviations and Numerical Equivalents Used in Tables

For Relative Abundance

a (abundant) indicates over 50 individuals oc (occasional) from 11 to 25 individuals

c (common) from 25 to 50 individuals r (rare) from 6 to 10 individuals

For Apparatus
n = sample taken by net p = sample taken by pump

Values enclosed in parentheses are interpolated

Table 1. Distributional and environmental records for Ceratium praelongum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
VC)

Salinity
(o/oo)

pH

P04
mg/m3

1

1

r

lb

7

50

r

15

101

50

r

IS

102

100

r

16

104

50

r

16

108

100

r

19

121

50

r

31

. 191

r

33

198A

50

r

47

267

50

r

47

268

100

r

47

269

r

48

273

100

r

49

278

50

r

55

327

50

r

56

329

50

r

56

330

100

r

69

424

r

76

471

50

r

77

472

50

oc

81

491

r

81

492

50

r

81

493

r

85

511

50

r

87

522

50

r

95

568A

r

96

569

50

r

96

572

50

r

97

575

50

r

98

581

r

98

583

100

r

98

584

r

98

588

0-100

oc

99

590

50

oc

99

591

100

oc

100

596

50

oc

100

597

100

oc

100

599

50

r

100

602

100

r

101

604

50

r

102

610

50

r

102

611

100

r

109

656

100

r

112

675

50

oc

113

681

50

r

113

684

50

r

114

687

50

r

132

803

100

r

152

945

50

r

152

949

50

r

153

955

r

153

956

50

r

154

958

r

155

965

oc

155

966

50

r

155

967

100

oc

156

972

oc

156

973

50

r

157

978

oc

157

979

50

oc

157

980

100

oc

158

983

oc

158

984

50

oc

158

985

100

oc

159

990

r

159

991

50

r

160

1003

100

r

n
n
n
n

P
n

P
n

P
n
n

P
n

n

P
n
n
n
P
P
n
n

P
n
n
n
n

P
n
n
n
P
P
n
n
n
n
P
P
n
n
n
n
n
n

P
n

P
n

P
P
P
n
n
n
n
n
n
n
n
n
n
n
n
n
n
n

5/12/28

5/16/28

8/11/28

8/11/28

8/13/28

8/13/28

8/20/28

10/ 3/28

10/ 8/28

11/23/28

11/23/28

11/23/28

11/25/28

11/27/28

12/16/28

12/18/28

12/18/28

1/12/29

2/16/29

2/18/29

2/26/29

2/26/29

2/26/29

3/ 6/29

3/11/29

4/25/29

4/26/29

4/26/29

4/28/29

4/30/29

4/30/29

4/30/29

4/30/29

5/ 2/29

5/ 2/29

5/ 4/29

5/ 4/29

5/ 4/29

5/ 4/29

5/ 7/29

5/ 9/29

5/ 9/29

5/29/29

6/ 5/29

6/25/29

6/25/29

6/27/29

9/ 8/29

10/27/29

10/27/29

10/29/29

10/29/29

10/31/29

11/ 2/29

11/ 2/29

11/ 2/29

11/ 4/29

11/ 4/29

11/ 6/29

11/ 6/29

11/ 6/29

11/ 8/29

11/ 8/29

11/ 8/29

11/11/29

11/11/29

11/13/29

24.0
(22.2
19.8
18.4
24.4
19.9
25.2
28.5
28.2
23.8
22.7
23.9
22.7
22.6
18.7
18.5
16.6
21.1
22.1
23.5
26.5
26.4
26.5
27.8
26.5
(29.4
29.2
29.2
28.0
27.0
26.7
27.0
26.7
27.8
27.8
27.6
27.6
27.6
27.6
26.2
25.8
25.6
19.4
21.7
23.8
23.8
16.2
14.3
14.2
14.2
28.1
28.1
28.3
27.8
27.7
27.2
27.6
27.0
27.1
27.1
26.8
28.2
28.2
27.6
28.6
28.5
28.5

36.2

8.16

34

36.3

8.21

38)

36.5

8.21

8

36.4

8.20

19

36.4

8.23

8

36.5

8.17

13

37.1

8.27

5

34.4

8.27

2

36.2

8.24

4

36.0

8.23

20

36.2

8.23

20

36.0

8.23

17

36.3

8.26

16

36.1

8.26

13

35.0

8.18

12

35.1

8.14

9

34.8

8.11

12

35.2

8.12

62

35.9

8.14

42

36.0

8.19

16

35.8

8.19

38

35.9

8.19

38

35.8

8.19

38

36.2

8.22

40

36.1

8.26

20

34.7

8.26

14)

35.3

8.23

12

35.3

8.23

12

35.4

8.16

21

35.3

8.16

24

35.4

8.14

32

35.3

8.16

24

35.4

8.14

32

34.9

8.22

12

35.0

8.22

17

34.7

8.21

10

34.7

8.22

12

34.7

8.21

10

34.7

8.22

12

34.7

8.24

8

35.0

8.24

8

35.0

8.23

8

34.8

8.18

5

34.6

8.23

7

34.6

8.25

5

34.6

8.25

5

34.6

8.04

63

33.4

8.30

16

34.5

7.87

53

34.5

7.87

53

34.2

8.47

7

34.4

8.39

7

34.2

8.39

7

34.9

8.29

29

34.9

8.30

30

35.0

8.30

35

35.0

8.34

28

35.1

8.37

46

35.3

8.27

47

35.2

8.32

60

35.5

8.30

64

35.6

8.34

36

35.6

8.39

50

35.9

8.39

48

35.7

8.37

15

35.7

8.39

15

35.7

8.44

16

49

50

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 2. Distributional and environmental records for Ceratium cephalotum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4

mg/m^

lb

8

100

r

n

5/16/28

(20.3

36.5

8.18

36)

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

48

273

100

r

n

11/25/28

22.7

36.3

8.26

16

83

501

r

n

3/ 2/29

27.5

36.3

8.24

29

89

529

50

r

n

3/23/29

28.6

35.8

8.27

12

90

533

r

n

3/25/29

28.5

35.5

8.27

21

90

534

50

oc

n

3/25/29

28.6

35.6

8.26

21

91

540

oc

n

3/27/29

28.7

35.1

8.30

21

91

541

50

oc

n

3/27/29

28.5

35.2

8.30

24

92

545

r

n

3/29/29

28.5

35.3

8.29

28

93

551

50

r

n

3/31/29

28.5

34.8

8.30

28

95

563

50

r

n

4/24/29

29.3

34.9

8.24

16

96

569

50

r

n

4/26/29

29.2

35.3

8.23

12

101

605

100

r

n

5/ 7/29

25.2

35.1

8.23

8

103

615

r

n

5/11/29

26.0

35.0

8.25

5

103

616

50

r

n

5/11/29

25.8

35.2

8.25

5

103

617

100

r

n

5/11/29

24.8

35.2

8.25

5

104

623

100

r

n

5/13/29

25.3

35.3

8.21

7

105

628

50

r

n

5/15/29

26.8

34.9

8.23

5

108

648

100

r

n

5/27/29

25.2

35.0

8.23

4

109

656

100

r

n

5/29/29

19.4

34.8

8.18

5

113

681

50

r

n

6/25/29

23.8

34.6

8.25

5

132

797

r

n

9/ 8/29

21.0

33.9

8.34

15

133

804

r

n

9/10/29

22.7

34.7

8.47

7

134

808

r

n

9/12/29

22.9

34.7

8.34

6

135

826

100

r

p

9/14/29

18.7

34.8

8.34

5

136

827

r

n

9/16/29

24.6

35.4

8.37

3

136

828

50

r

n

9/16/29

21.4

35.1

8.39

3

140

860

r

p

10/ 3/29

26.9

35.0

8.42

7

141

864

r

n

10/ 5/29

25.9

35.2

8.34

5

141

865

50

r

n

10/ 5/29

24.8

35.3

8.34

5

142

874

100

r

n

10/ 7/29

16.6

34.4

8.27

7

142

876

r

P

10/ 7/29

24.1

34.8

8.33

5

143

883

100

r

n

10/ 9/29

13.8

34.1

8.30

10

144

886

r

n

10/11/29

23.3

35.0

8.37

6

145

892

100

r

n

10/13/29

16.0

34.1

8.31

6

146

898

100

r

n

10/15/29

19.7

34.3

8.26

7

147

905

100

r

n

10/17/29

19.2

35.0

8.29

5

150

931

100

r

n

10/23/29

19.6

34.6

8.32

U

Table 3. Distributional and environmental records for Ceratium gravidum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(0/00)

pH

PO4

mg/m3

1

1

r

n

5/12/28

24.0

36.2

8.16

34

lb

6

r

n

5/16/28

(22.3

36.3

8.20

46)

lb

7

50

oc

n

5/16/28

122.2

36.4

8.21

39)

lb

8

100

oc

n

5/16/28

120.3

35.5

8.18

36)

15

98

50

r

p

8/11/28

19.8

36.5

8.21

8

15

99

100

r

p

8/11/28

18.4

36.4

8.20

19

15

101

50

oc

n

8/11/28

19.8

36.5

8.21

8

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

18

116

100

r

p

8/17/28

20.4

36.8

8.21

5

18

119

100

oc

n

8/17/28

20.4

36.8

8.21

5

19

124

100

oc

n

8/20/28

22.4

37.0

8.25

5

20

129

50

r

n

8/22/28

25.8

36.6

8.26

3

20

130

100

oc

n

8/22/28

22.6

36.7

8.19

5

22

143

50

r

n

8/27/28

24.5

36.2

8.21

9

23

149

50

r

n

8/29/28

20.9

36.0

8.14

13

24

155

50

oc

n

8/31/28

23.1

36.0

8.14

8

25

158

50

r

p

9/ 3/28

21.5

36.0

8.22

12

25

161

50

oc

n

9/ 3/28

21.5

36.0

8.22

12

25

161

50

r

n

9/ 3/28

21.5

36.0

8.22

12

25

162

100

r

n

9/ 3/28

14.6

35.7

7.93

121

26

164

50

r

p

9/ 5/28

24.1

36.1

8.21

5

27

168

50

r

p

9/ 7/28

26.0

36.2

8.30

4

27

171

50

oc

n

9/ 7/28

26.0

36.2

8.30

4

27

171

50

oc

n

9/ 7/28

26.0

36.2

8.30

4

28

177

50

r

n

S/11/28

26.7

36.3

8.26

4

SO

189

50

r

n

9/15/28

27.8

36.1

8.29

3

31

192

50

r

n

10/ 3/28

28.2

35.4

8.23

2

APPENDIX

51

Table 4. Distributional and environmental records for Ceratium digitatum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

PO4

mg/m3

25

158

50

r

p

9/ 3/28

21.5

36.0

8.22

12

26

164

50

r

p

9/ 5/28

24.1

36.1

8.21

5

45

257

50

r

n

11/19/28

22.4

35.2

8.13

46

46

265

50

r

p

11/21/28

23.2

35.3

8.16

40

49

279

100

r

n

11/27/28

21.6

35.9

8.26

13

49a

283

r

n

11/28/28

(23.2

36.0

8.23

13)

SO

287

r

P

11/29/28

23.2

36.0

8.23

13

52

302

r

p

12/ 3/28

22.5

35.4

8.21

8

53

305

100

r

n

12/ 5/28

IS. 9

35.6

8.19

13

54

322

r

p

12/14/28

23.4

35.5

8.22

9

78

478

r

p

2/20/29

24.6

36.0

8.17

32

83

502

50

r

n

3/ 2/29

27.4

36.5

8.24

25

92

546

50

r

n

3/29/29

28.4

35.4

8.29

28

95

564

100

r

n

4/24/29

28.5

35.4

8.22

21

99

591

100

r

n

5/ 2/29

27.8

35.0

8.22

17

100

596

50

r

n

5/ 4/29

27.6

34.7

8.21

10

102

611

100

r

n

5/ 9/29

25.6

35.0

8.23

8

102

612

r

p

5/ 9/29

25.8

35.0

8.24

8

105

629

100

r

n

5/15/29

25.2

35.1

8.23

5

107

641

100

r

n

5/19/29

26.8

34.9

8.23

11

136

829

100

r

n

9/16/29

18.6

35.0

8.39

3

140

858

100

r

n

10/ 3/29

25.5

35.0

8.34

7

145

891

50

r

n

10/13/29

18.7

34.3

8.34

6

147

905

100

r

n

10/17/29

19.2

35.0

8.29

5

151

939

100

r

n

10/26/29

12.5

34.6

155

967

100

r

n

11/ 2/29

27.2

35.0

a.'so

35

Table 5. Distributional and environmental records for Ceratium candelabrxim

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/m3

1

1

oc

n

5/12/28

24.0

36.2

8.16

34

la

4

r

n

5/14/28

(22.3

36.3

8.20

46)

la

5

r

n

5/14/28

22.3

36.3

8.20

46'i

lb

6

oc

n

5/16/28

(22.3

36.3

8.20

46)

lb

7

so

oc

n

5/16/28

22.2

36.3

8.20

39)

2

14

r

n

5/18/28

20.5

36.4

8.23

58

13a

89

r

n

8/ 8/28

(16.3

34.0

8.14

15)

14

92

50

r

p

8/ 9/28

15.0

35.1

8.18

16

14

93

r

p

8/ 9/28

21.2

35.2

8.18

11

14

95

50

r

n

8/ 9/28

15.0

35.1

8.18

16

IS

99

100

r

P

8/11/28

18.4

36.4

8.20

19

15

102

100

oc

n

8/11/28

18.4

36.4

8.20

19

16

103

r

p

8/13/28

25.9

36.2

8.24

8

16

104

50

r

P

8/13/28

24.4

36.4

8.23

8

16

106

r

n

8/13/28

25.9

36.2

8.24

8

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

17

109

oc

p

8/15/28

26.2

36.6

8.29

9

17

111

r

n

8/15/28

26.2

36.6

8.29

9

17

112

50

r

n

8/15/28

21.9

36.6

8.28

12

18

114

r

p

8/17/28

27.0

37.0

8.23

5

18

115

50

oc

p

8/17/28

22.4

36.8

8.24

5

18

116

100

r

p

8/17/28

20.4

36.8

8.21

5

18

117

oc

n

8/17/28

27.0

37.0

8.23

5

18

118

SO

r

n

8/17/28

22.4

36.8

8.24

5

19

123

50

r

n

8/20/28

25.2

37.1

8.27

5

19

124

100

r

n

8/20/28

22.4

37.0

8.25

5

20

126

50

r

p

8/22/28

25.8

36.6

8.26

3

20

128

r

n

8/22/28

26.0

36.6

8.37

5

20

129

SO

oc

n

8/22/28

25.8

36.6

8.26

3

20

130

100

oc

n

8/22/28

22.6

36.7

8.19

5

20-21

131

c

n

8/23/28

(26.6

36.3

8.32

4)

21

134

100

c

p

8/24/28

21.0

36.8

8.20

7

21

135

oc

n

8/24/28

26.6

36.3

8.32

4

21

137

100

r

n

8/24/28

21.0

36.8

8.20

7

22

139

oc

P

8/27/28

26.7

36.0

8.26

8

22

143

so

r

n

8/27/28

24.5

36.2

8.21

9

23

145

oc

P

8/27/28

26.7

36.0

8.26

8

23

149

so

oc

n

8/27/28

24.5

36.2

8.21

9

23

150

100

r

n

8/29/28

16.6

36.0

8.18

75

24

152

50

r

P

8/31/28

23.1

36.0

8.14

8

25

157

oc

P

9/ 3/28

27.5

35.6

8.31

5

52

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 5. Distributional and environmental records for Ceratium candelabrum- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

CC)

(o/oo)

mg/m3

25

158

50

oc

P

9/ 3/28

21.5

36.0

8.22

12

25

160

r

n

9/ 3/28

27.5

35.6

8.31

5

25

161

50

oc

n

9/ 3/28

21.5

36.0

8.22

12

26

164

50

r

p

9/ 5/28

24.1

36.1

8.21

5

26

165

100

r

p

9/ 5/28

14.9

35.6

8.11

40

27

171

50

r

n

9/ 7/28

26.0

36.2

8.30

4

28

177

50

r

n

9/11/28

26.7

36.3

8.26

4

30

189

50

r

n

9/15/28

27.8

36.1

8.29

3

31

193

100

r

n

10/ 3/28

23.4

36.5

8.19

28

32

194A

r

p

10/ 5/28

28.0

36.0

8.23

2

32

195

50

r

n

10/ 5/28

27.2

36.0

8.24

2

32

196

100

r

n

10/ 5/28

22.2

36.4

8.10

30

33

197 A

r

p

10/ 8/28

28.5

35.6

8.23

4

34

200

oc

n

10/ 9/28

28.5

35.9

8.28

2

34

200A

r

p

10/ 9/28

28.5

35.9

8.28

2

34

201

100

oc

n

10/ 9/28

20.5

36.6

8.16

16

35

205

100

r

n

10/26/28

14.4

34.9

7.88

189

35

206

r

p

10/26/28

27.4

29.7

8.31

15

35-36

209

r

n

10/27/28

(27.0
(27.0

30.0

8.27

16)

35-36

211

r

n

10/27/28

30.0

8.27

16)

36

213

r

n

10/30/28

26.5

31.5

8.23

16

36

215

100

r

n

10/30/28

14.4

34.9

7.85

149

36

216

r

P

10/30/28

26.5

31.5

8.23

16

37

218

50

r

n

11/ 1/28

18.8

34.5

8.00

121

37

219

100

r

n

11/ 1/28

15.1

34.9

7.82

153

40

232

r

P

11/ 8/28

22.2

33.7

8.21

24

41

238

r

P

11/10/28

20.4

34.2

8.11

32

41a

240

r

n

11/12/28

(19.6

34.5

8.09

39)

43

244

r

n

11/15/28

19.6

34.8

8.09

52

44

250

oc

n

11/17/28

20.7

34.9

8.03

38

44

251

50

oc

n

11/17/28

20.4

34.9

8.04

34

44

253

oc

p

11/17/28

20.7

34.9

8.03

38

45

256

oc

n

11/19/28

22.4

35.3

8.12

38

45

257

50

oc

n

11/19/28

22.4

35.2

8.13

46

45

258

100

r

n

11/19/28

18.6

35.1

8.00

50

45

259

r

P

1-1/19/28

22.4

35.3

8.12

38

45

260

50

r

p

11/19/28

22.4

35.2

8.13

46

46

261

r

n

11/21/28

23.3

35.3

8.16

36

46

262

50

r

n

11/21/28

23.2

35.3

8.16

40

46

263

100

oc

n

11/21/28

22.5

35.4

8.17

40

46

264

oc

p

11/21/28

23.3

35.3

8.16

36

47

266

r

n

11/23/28

23.9

36.0

8.23

17

47

267

50

oc

n

11/23/28

23.8

36.0

8.23

20

47

268

100

oc

n

11/23/28

22.7

36.2

8.23

20

47

269

oc

p

11/23/28

23.9

36.0

8.23

17

48

273

100

oc

n

11/25/28

22.7

36.3

8.26

16

48

274

r

p

11/25/28

23.6

36.4

8.23

13

49

278

50

r

n

11/27/28

22.6

36.1

8.26

13

49

279

100

r

n

11/27/28

21.6

35.9

8.26

13

50

285

50

r

n

11/29/28

22.0

35.9

8.23

13

50

288

50

oc

p

11/29/28

22.0

35.9

8.23

13

51

294

50

r

n

12/ 1/28

20.5

35.6

8.22

17

51

295

100

r

n

12/ 1/28

20.0

35.6

8.22

17

52

300

50

oc

n

12/ 3/28

20.2

35.6

8.20

8

52

301

100

oc

n

12/ 3/28

18.2

35.2

8.17

8

53

305

100

r

n

12/ 5/28

19.9

35.6

8.19

13

54

320

50

oc

n

12/14/28

19.8

35.4

8.18

17

54

321

100

oc

n

12/14/28

18.7

35.4

8.16

20

55

324

50

oc

n

12/16/28

18.7

35.0

8.18

12

55

325

100

oc

n

12/16/28

16.7

34.9

7.17

12

56

328

oc

n

12/18/28

20.8

34.9

8.13

9

56

329

50

oc

n

12/18/28

18.5

35.1

8.14

9

56

330

100

oc

n

12/18/28

16.6

34.8

8.11

12

56

332

50

r

p

12/18/28

18.5

35.1

8.14

9

56

333

100

r

P

12/18/28

16.6

34.8

8.11

12

56-57

334

oc

n

12/19/28

(19.0

34.5

8.14

20)

57

335

r

n

12/20/28

19.0

34.5

8.14

20

57

336

50

oc

n

12/20/28

15.6

34.3

8.14

21

57

337

100

r

n

12/20/28

14.3

34.4

8.10

40

57

338

r

P

12/20/28

19.0

34.5

8.14

20

57

340

100

r

P

12/20/28

14.3

34.4

8.10

40

58

341

r

n

12/22/28

17.0

34.0

8.12

20

58

342

50

r

n

12/22/28

14.8

34.0

8.12

25

59

351

r

p

12/24/28

16.3

34.0

8.10

38

61

363

oc

n

12/28/28

16.9

34.0

8.05

46

61

364

50

r

n

12/28/28

14.0

34.0

8.05

60

APPENDIX

S3

Table 5. Distributional and environmental records for Ceratlum candelabrum --Continued

Depth

Relative

Appa-

Temperature

Salinity

pH

P04

Station

Sample

(m)

abundance

ratus

Date

("C)

(o/oo)

mg/m3

61

365

100

r

n

12/28/28

10.8

34.0

8.03

80

61

366

r

P

12/28/28

16.9

34.0

8.05

46

61-62

369

oc

n

12/29/28

(16.9
(16.9

34.0

8.05

46)
46)

61-62

370

c

n

12/29/28

34.0

8.05

62

371

c

n

12/30/28

19.2

34.2

8.12

32

62

372

50

c

n

12/30/28

16.2

34.3

8.10

28

62

373

100

c

n

12/30/28

13.1

34.2

8.06

48

62

374

oc

P

12/30/28

19.2

34.2

8.12

32

62

375

50

r

P

12/30/28

16.2

34.3

8.10

28

63

379

r

n

1/ 1/29

20.5

34.6

8.07

21

63

381

100

r

n

1/ 1/29

15.6

34.6

8.08

24

63

384

100

r

P

1/ 1/29

15.6

34.6

8.08

24

63-64

387

r

n

1/ 2/29

20.5

34.6

8.07

21i
21)

63-64

390

r

n

1/ 2/29

20.5

34.6

8.07

64

395

50

r

n

1/ 3/29

17.2

34.6

8.12

29

64

396

100

r

n

1/ 3/29

15.8

34.5

8.10

32

64-65

401

oc

n

1/ 3/29

(20.6

34.6

8.12

21)

64-65

402

oc

n

1/ 4/29

(20.6

34.6

8.12

21

64-65

403

r

n

1/ 4/29

20.6

34.6

8.12

21)

64-65

404

r

n

1/ 4/29

(20.6

34.6

8.12

2l)

65

408

r

p

1/ 5/29

20.2

34.5

8.10

24

65

409

50

r

P

1/ 5/29

16.5

34.5

8.10

25

66

415

100

r

p

1/ 7/29

17.8

34.9

8.12

21

67

416

r

n

1/ 8/29

19.3

34.9

8.11

21

67

418

100

oc

n

1/ 8/29

16.2

34.6

8.05

40

73

452

100

r

n

2/10/29

14.7

35.0

7.80

178

75

462

oc

n

2/14/29

22.8

35.8

8.18

44

76

467

oc

n

2/16/29

23.4

35.9

8.15

50

76

468

50

r

n

2/16/29

22.1

35.9

8.14

42

76

469

100

r

n

2/16/29

21.2

35.8

8.12

45

76

471

50

r

p

2/16/29

22.1

35.9

8.14

42

77

474

50

oc

P

2/18/29

23.5

36.0

8.19

16

78

475

oc

n

2/20/29

24.6

36.0

8.17

32

78

476

50

oc

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

478

oc

p

2/20/29

24.6

36.0

8.17

32

78

479

50

oc

p

2/20/29

23.8

36.1

8.14

32

79

481

oc

n

2/22/29

25.2

36.0

8.17

34

79

482

50

oc

n

2/22/29

24.5

36.1

8.17

34

79

483

100

oc

n

2/22/29

21.8

36.2

8.13

45

79

484

oc

P

2/22/29

25.2

36.0

8.17

34

79

485

50

oc

p

2/22/29

24.5

36.1

8.17

34

80

486

oc

n

2/24/29

26.0

35.9

8.20

36

80

487

50

r

n

2/24/29

25.9

36.0

8.19

29

80

488

oc

p

2/24/29

26.0

35.9

8.20

36

80

489

50

oc

p

2/24/29

25.9

36.0

8.19

29

80

490

100

r

p

2/24/29

23.4

36.2

8.16

32

81

491

c

n

2/26/29

26.5

35.8

8.19

38

81

492

50

oc

n

2/26/29

26.4

35.9

8.19

38

81

493

r

p

2/26/29

26.5

35.8

8.19

38

81

494

50

r

p

2/26/29

26.4

35.9

8.19

38

31

495

100

r

p

2/26/29

23.6

36.2

8.18

36

)2

496

oc

n

2/28/29

27.2

36.3

8.21

34

)2

497

50

r

n

2/28/29

27.2

36.3

8.21

34

83

502

50

r

n

3/ 2/29

27.4

36.5

8.24

25

84

506

50

oc

n

3/ 4/29

27.5

36.4

8.21

24

85

510

oc

n

3/ 6/29

27.9

36.2

8.22

40

85

511

50

r

n

3/ 6/29

27.8

36.2

8.22

40

85

513

50

r

p

3/ 6/29

27.8

36.2

8.22

40

86

517

50

r

n

3/ 9/29

27.4

36.2

8.29

17

86

519

50

r

p

3/ 9/29

27.4

36.2

8.29

17

87

522

50

oc

n

3/11/29

26.5

36.1

8.26

20

87

524

50

r

p

3/11/29

26.5

36.1

8.26

20

88

526

oc

n

3/21/29

28.5

35.9

8.23

16

88

527

50

oc

n

3/21/29

28.4

35.9

8.25

13

89

528

oc

n

3/23/29

28.4

35.6

8.25

21

89

529

50

oc

n

3/23/29

28.6

35.8

8.27

12

90

533

c

n

3/25/29

28.5

35.5

8.27

21

90

534

50

oc

n

3/25/29

28.6

35.6

8.26

21

90

535

r

p

3/25/29

28.5

35.5

8.27

21

90

536

50

r

p

3/25/29

28.6

35.6

8.26

21

90

537

100

r

p

3/25/29

26.3

35.8

8.24

21

91

540

r

n

3/27/29

28.7

35.1

8.30

21

91

541

50

c

n

3/27/29

28.5

35.2

8.30

24

91

542

r

p

3/27/29

28.7

35.1

8.30

21

91

543

50

r

P

3/27/29

28.5

35.2

8.30

24

54

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 5. Distributional and environmental records for Ceratium candelabrum --Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature

("C)

Salinity
(o/oo)

pH

PO4

mg/m3

91

544

100

oc

P

3/27/29

25.8

36.0

8.25

30

92

546

50

c

n

3/29/29

28.4

35.4

8.29

28

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

551

50

r

n

3/31/29

28.5

34.8

8.30

28

93

552

r

p

3/31/29

28.7

34.7

8.30

28

Samoa

555
558

50

r
r

n
n

29. Z

34! 7

8. 2 5

94

4/22/29

i'4

94

560

r

P

4/22/29

29.5

34.7

8.25

14

95

563

50

oc

n

4/24/29

29.3

34.9

8.24

16

95

564

100

r

n

4/24/29

28.5

35.4

8.22

21

95

565

r

P

4/24/29

29.4

34.7

8.26

14

96

568

oc

n

4/26/29

29.3

35.3

8.23

12

95

568A

r

n

4/25/29

(29.4

34.7

8.26

14)

96

569

50

r

n

4/26/29

29.2

35.3

8.23

12

96

572

50

r

P

4/26/29

29.2

35.3

8.23

12

96

573

100

r

P

4/26/29

28.2

35.7

8.19

25

97

574

oc

n

4/28/29

28.3

35.2

8.16

24

97

575

50

oc

n

4/28/29

28.0

35.4

8.16

21

97

576

100

oc

n

4/28/29

27.6

35.6

8.15

25

97

579

100

r

P

4/28/29

27.6

35.6

8.15

25

98

581

oc

n

4/30/29

27.0

35.3

8.16

24

98

588

100

r

P

4/30/29

26.7

35.4

8.14

32

99

590

50

c

n

5/ 2/29

27.8

34.9

8.22

12

99

591

100

c

n

5/ 2/29

27.8

35.0

8.22

17

99

592

oc

P

5/ 2/29

27.9

34.9

8.21

12

99

593

50

r

P

5/ 2/29

27.8

34.9

8.22

12

99

594

100

oc

P

5/ 2/29

27.8

35.0

8.22

17

99

595

oc

n

5/ 2/29

27.9

34.9

8.21

12

100

596

50

c

n

5/ 4/29

' 27.6

34.7

8.21

10

100

597

100

c

n

5/ 4/29

27.6

34.7

8.22

12

100

598

oc

P

5/ 4/29

27.7

34.7

8.21

10

100

599

50

oc

P

5/ 4/29

27.6

34.7

8.21

10

100

602

100

oc

P

5/ 4/29

27.6

34.7

8.22

12

101

604

50

oc

n

5/ 7/29

26.2

34.7

8.24

8

101

605

100

oc

n

5/ 7/29

25.2

35.1

8.23

8

101

606

r

P

5/ 7/29

26.3

34.7

8.24

8

102

609

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

612

r

P

5/ 9/29

25.8

35.0

8.24

8

102

613

50

r

p

5/ 9/29

25.8

35.0

8.24

8

103

615

r

n

5/11/29

26.0

35.0

8.25

5

103

616

50

r

n

5/11/29

25.8

35.2

8.25

5

103

617

100

oc

n

5/11/29

24.8

35.2

8.25

5

103

618

r

P

5/11/29

26.0

35.0

8.25

5

104

621

r

n

5/13/29

26.1

35.2

8.24

7

104

622

50

r

n

5/13/29

25.8

35.2

8.24

7

104

625

50

r

P

5/13/29

25.8

35.2

8.24

7

105

628

50

r

n

5/15/29

26.8

34.9

8.23

5

105

630

r

P

5/15/29

26.9

34.9

8.23

5

106

633

r

n

5/17/29

27.2

35.0

8.23

5

106

634

50

r

n

5/17/29

27.0

35.0

8.23

5

106

635

100

r

n

5/17/29

25.6

35.1

8.23

5

107

639

r

n

5/19/29

28.0

34.4

8.23

5

107

640

50

oc

n

5/19/29

27.9

34.4

8.23

4

107

641

100

r

n

5/19/29

26.8

34.9

8.23

11

107

642

r

P

5/19/29

28.0

34.4

8.23

5

107

645

50

r

P

5/19/29

27.9

34.4

8.23

4

108

646

r

n

5/27/29

28.4

35.0

8.25

4

108

647

50

r

n

5/27/29

26.8

35.0

8.24

4

108

648

100

r

n

5/27/29

25.2

35.0

8.23

4

109

655

50

oc

n

5/29/29

23.1

35.0

8.22

3

109

656

100

c

n

5/29/29

19.4

34.8

8.18

5

109

658

50

r

P

5/29/29

23.1

35.0

8.22

3

110

661

c

n

5/31/29

23.9

34.7

8.18

5

110

663

100

oc

n

5/31/29

17.9

34.7

8.14

11

110

664

oc

P

5/31/29

23.9

34.7

8.18

5

110

665

50

oc

P

5/31/29

18.4

34.8

8.16

7

111

668

oc

n

6/ 3/29

20.1

34.5

8.18

5

111

671

oc

P

6/ 3/29

20.1

34.5

8.18

5

112

675

50

r

n

6/ 5/29

21.7

34.6

8.23

7

112

676

100

r

n

6/ 5/29

19.8

34.7

8.20

8

112

677

r

P

6/ 5/29

23.2

34.6

8.22

7

113

680

oc

n

6/25/29

24.2

34.5

8.25

5

113

681

50

oc

n

6/25/29

23.8

34.6

8.25

5

113

682

100

r

n

6/25/29

21.5

34.7

8.23

8

113

684

50

r

P

6/25/29

23.8

34.6

8.25

5

114

686

oc

n

6/27/29

19.9

34.3

8.15

7

APPENDIX

55

Table 5. Distributional and environmental records for Ceratium candelabrum --Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

PO4

mg/m3

114

687

50

DC

n

114

688

100

oc

n

114

689

DC

P

lis

692

r

n

115

693

50

r

n

117

709

50

r

n

128

779

oc

n

129

780

r

n

130

781

oc

n

130

784

oc

p

130

785

50

r

P

130

786

100

r

P

130

787

c

n

131

789

100

oc

n

131

792

oc

P

131

793

r

P

132

797

oc

n

132

798

50

r

n

132

799

100

oc

n

132

801

r

P

132

802

50

r

p

132

803

100

r

P

133

805

50

r

n

134

808

oc

n

134

809

50

r

n

134

810

100

r

n

134

815

r

P

135

821

50

r

n

135

822

100

r

n

135

825

50

r

P

136

828

50

r

n

136

832

r

P

137

836

r

n

139

850

50

r

n

139

851

100

r

n

140

858

100

r

n

141

865

50

r

n

142

873

50

oc

n

142

874

100

oc

n

142

877

50

r

P

143

881

r

n

143

883

100

r

n

143

885

50

r

P

145

891

50

r

n

146

896

r

n

146

897

50

r

n

146

898

100

oc

n

147

904

50

r

n

149

921

50

r

n

149

927

r

P

150

929

r

n

150

930

50

oc

n

150

931

100

oc

n

151

937

oc

n

151

938

50

oc

n

151

939

100

n

151

941

P

152

944

oc

n

152

945

50

oc

n

152

946

100

n

152

948

P

153

951

oc

n

153

952

50

n

153

953

100

oc

n

154

959

50

n

155

965

n

155

966

50

n

155

967

100

oc

n

156

972

n

156

973

50

n

156

974

100

oc

n

157

978

oc

n

157

979

50

n

157

980

lOU

n

158

983

c

n

158

984

50

oc

n

6/27/29

6/27/29

6/27/29

6/29/29

6/29/29

7/ 3/29

7/25/29

7/27/29

9/ 4/29

9/ 4/29

9/ 4/29

9/ 4/29

9/ 4/29

9/ 6/29

9/ 4/29

9/ 6/29

9/ 8/29

9/ 8/29

9/ 8/29

9/ 8/29

9/ 8/29

9/ 8/29

9/10/29

9/12/29

9/12/29

9/12/29

9/12/29

9/14/29

9/14/29

9/14/29

9/16/29

9/16/29

9/18/29

9/22/29

9/22/29

10/ 3/29

10/ 5/29

10/ 7/29

10/ 7/29

10/ 7/29

10/ 9/29

10/ 9/29

10/ 9/29

10/13/29

10/15/29

10/15/29

10/15/29

10/17/29

10/21/29

10/21/29

10/23/29

10/23/29

10/23/29

10/26/29

10/26/29

10/26/29

10/26/29

10/27/29

10/27/29

10/27/29

10/27/29

10/29/29

10/29/29

10/29/29

10/31/29

11/ 2/29

11/ 2/29

11/ 2/29

11/ 4/29

11/ 4/29

11/ 4/29

11/ 6/29

11/ 6/29

11/ 6/29

11/ 8/29

11/ 8/29

16.2
13.0
19.9
20.6
17.5
12.5
16.4
16.3
16.2
16.2
11.7
8.8
16.2
12.1
16.2
19.3
21.0
17.6
14.3
21.0
17.6
14.3
20.8
22.9
19.8
18.1
22.9
21.5
18.7
21.5
21.4
24.6
25.5
25.8
22.4
25.5
24.8
21.8
16.6
21.8
22.4
13.8
19.0
18.7
22.4
22.4
19.7
23.1
23.3
23.5
25.6
22.8
19.6
26.0
18.3
12.5
26.0
27.4
14.2
11.4
27.4
28.1
28.1
20.5
28.2
27.8
27.7
27.2
27.6
27.0
26.4
27.1
27.1
26.8
28.2
28.2

34.6

8.04

63

34.5

8.00

91

34.3

8.15

7

34.6

8.19

4

34.6

8.12

17

34.2

8.06

51

33.0

8.12

29

33.1

8.13

25

33.4

8.34

36

33.4

8.34

36

33.4

8.26

83

33.7

8.06

176

33.4

8.34

36

33.4

8.32

33.4

8.34

36

33.4

8.34

• ••

33.9

8.34

15

33.9

8.33

19

33.4

8.30

16

33.9

8.34

15

33.9

8.33

19

33.4

8.30

16

34.7

8.37

7

34.7

8.34

6

34.6

8.34

6

34.6

8.34

6

34.7

8.34

6

35.0

8.37

5

34.8

8.34

5

35.0

8.37

5

35.1

8.39

3

35.4

8.37

3

35.0

8.39

4

34.9

8.31

6

35.2

8.28

6

35.0

8.34

7

35.3

8.34

5

34.8

8.30

5

34.4

8.27

7

34.8

8.30

5

34.4

8.30

6

34.1

8.30

10

34.2

8.34

6

34.3

8.34

6

34.9

8.37

6

34.9

8.30

6

34.3

8.26

7

35.3

8.29

5

35.0

8.37

6

35.0

8.34

6

34.7

8.39

7

34.8

8.35

10

34.6

8.32

11

34.0

34.4

• • •

34.6

...

34.0

• . >

33.7

8.35

20

34.5

7.87

53

34.7

7.76

75

33.7

8.35

20

34.2

8.47

7

34.4

8.39

7

34.7

8.28

31

34.2

8.40

7

34.9

8.29

29

34.9

8.30

30

35.0

8.30

35

35.0

8.34

28

35.1

8.37

46

35.1

8.30

48

35.3

8.27

47

35.2

8.32

60

35.5

8.30

64

35.6

8.34

36

35.6

8.39

50

56

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 5. Distributional and environmental records for Ceratium candelabrum — Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
("C)

Salinity
(o/oo)

pH

PO4
mg/m3

158

985

100

c

n

11/ 8/29

27.6

35.9

8.39

48

159

990

oc

n

11/11/29

28.6

35.7

8.37

15

159

991

50

r

n

11/11/29

28.5

35.7

8.39

15

159

992

100

r

n

11/11/29

28.0

35.7

8.37

23

160

1000

r

n

11/13/29

28.6

35.6

8.37

12

160

1002

50

oc

n

11/13/29

28.6

35.6

8.39

15

160

1003

100

r

n

11/13/29

28.5

35.7

8.44

16

Table 6. Distributional and environmental records for Ceratium furca Ehrenbergi

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4
mg/m3

(m)

abundance

ratus

VC)

(0/00)

5a

33

c

n

5/30/28

(12.4
112.4

35.6

8.15

21)
21)

5a

34

r

n

5/30/28

35.6

8.15

6

35

c

n

5/31/28

12.4

35.6

8.15

21

6

36

50

c

n

5/31/28

11.6

35.6

8.12

32

6

37

100

c

n

5/31/28

11.3

35.6

8.08

41

6a

42

c

n

5/31/28

(12.4

35.6

8.15

21

6b

43

c

n

6/ 2/28

12.4

35.6

8.15

21

6b

44

oc

n

6/ 2/28

(12.4

35.6

8.15

21

6b

45

oc

n

6/ 2/28

(12.4

35.6

8.15

21

6b

46

c

n

6/ 2/28

12.4

35.6

8.15

21

6c

47

100

c

n

6/ 2/28

(11.3

35.6

8.08

41

6c

48

oc

n

6/ 3/28

(12.4

35.6

8.15

21

6c

49

c

n

6/ 4/28

(12.4

35.6

8.15

21

6c

52

r

n

7/ 8/28

10.6

35.4

8.12

27

6g

54

c

n

7/10/28

10.6

35.4

8.12

27

6h

55

a

n

7/11/28

(10.6

35.4

8.12

27

7

57

50

oc

n

7/13/28

8.2

35.2

8.03

47

7

58

100

r

n

7/13/28

8.1

35.2

8.04

57

8

59

r

n

7/15/28

10.3

35.2

7.93

13

8

60

50

r

n

7/15/28

9.1

35.2

7.95

27

9

64

r

n

7/28/28

11.2

35.1

8.08

20

9

65

50

oc

n

7/28/28

8.4

35.1

7.96

55

9

66

100

oc

n

7/28/28

7.6

35.1

7.98

56

10

68

oc

n

7/30/28

10.9

34.9

8.08

28

10

69

50

r

n

7/30/28

10.0

34.9

8.04

34

10

70

100

oc

n

7/30/28

6.6

35.0

7.95

52

11a

78

oc

n

8/ 2/28

(10.7
(21.2

34.9

8.06

27)
11)

13a

89

r

n

8/ 2/28

35.2

8.18

14

93

r

p

8/ 9/28

21.2

35.2

8.18

11

15

97

r

p

8/11/28

24.8

36.4

8.21

11

16

105

100

r

p

8/13/28

19.9

36.3

8.17

13

16

108

100

r

n

8/13/28

19.9

36.3

8.17

13

17

111

r

n

8/15/28

26.2

36.6

8.29

9

19

122

r

n

8/20/28

26.6

37.0

8.34

5

20-21

131

r

n

8/23/28

(26.6

36.3

8.32

4)

21

134

100

r

p

8/24/28

21.0

36.8

8.20

7

23

145

oc

P

8/29/28

27.2

35.9

8.25

4

24

151

r

P

8/31/28

27.2

35.2

8.32

4

24

156

100

r

n

8/31/28

15.6

35.6

7.96

99

32

194A

r

p

10/ 5/28

28.0

36.0

8.23

2

40

232

r

P

11/ 8/28

22.2

33.7

8.21

24

40

233

50

r

p

11/ 8/28

15.3

34.9

7.87

161

41a

240

r

n

11/12/28

(18.7

34.7

8.06

45)

42

241

r

n

11/13/28

18.7

34.7

8.06

45

42

242

50

r

n

11/13/28

17.2

34.9

7.99

68

42

243

100

r

n

11/13/28

13.8

35.0

7.91

150

43

245

50

oc

n

11/15/28

17.0

34.9

7.93

80

43

246

100

oc

n

11/15/28

13.6

35.0

7.90

92

43

247

r

P

11/15/28

19.6

34.8

8.09

52

43

249

100

r

P

11/15/28

13.6

35.0

7.90

92

44'

250

r

n

11/17/28

20.7

34.9

8.03

38

44

251

50

r

n

11/17/28

20.4

34.9

8.04

34

44

252

100

r

n

11/17/28

13.8

35.0

7.85

70

44

253

oc

P

11/17/28

20.7

34.9

8.03

38

44

254

50

r

P

11/17/28

20.4

34.9

8.04

34

45

257

50

r

n

11/19/28

22.4

35.2

8.13

46

4S

258

100

r

n

11/19/28

18.6

35.1

8.00

50

45

259

r

P

11/19/28

22.4

35.3

8.12

38

45

260

50-0

r

P

11/19/28

22.4

35.2

8.13

46

46

263

100

oc

n

11/21/28

22.5

35.4

8.17

40

APPENDIX

57

Table 6. Distributional and environmental records for Ceratium furca Ehrenbergi--Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
("C)

Salinity
(o/oo)

pH

PO4

mg/m3

46

264

oc

P

11/21/28

23.3

35.3

8.16

36

47

266

r

n

11/23/28

23.9

36.0

8.23

17

47

267

50

r

n

11/23/28

23.8

36.0

8.23

20

47

268

100

oc

n

11/23/28

22.7

36.2

8.23

20

47

269

r

P

11/23/28

23.9

36.0

8.23

17

47

270

50

r

P

11/23/28

23.8

36.0

8.23

20

50

289

100

r

P

11/29/28

20.5

35.7

8.22

13

56-57

334

r

n

12/19/28

(19.0

34.5

8.14

20)

57

336

50

r

n

12/20/28

15.6

34.3

8.14

21

58

341

oc

n

12/22/28

17.0

34.0

8.12

20

58

342

50

c

n

12/22/28

14.8

34.0

8.12

25

58

344

oc

P

12/22/28

17.0

34.0

8.12

20

58

345

50

oc

P

12/22/28

14.8

34.0

8.12

25

58

346

100

oc

P

12/22/28

12.3

34.1

8.05

40

59

347

oc

n

12/24/28

16.3

34.0

8.10

38

59

348

oc

n

12/24/28

16.3

34.0

8.10

38

59

349

50

r

n

12/24/28

14.0

34.0

8.08

38

59

350

100

c

n

12/24/28

11.4

34.1

8.03

72

59

351

c

P

12/24/28

16.3

34.0

8.10

38

59

352

50

r

P

12/24/28

14.0

34.0

8.08

38

59

353

100

r

P

12/24/28

11.4

34.1

8.03

72

60

354

oc

n

12/26/28

15.0

34.0

8.07

50

60

355

50

oc

n

12/26/28

13.4

34.0

8.06

54

60

356

100

oc

n

12/26/28

10.6

34.0

8.03

62

60

357

oc

P

12/26/28

15.0

34.0

8.07

50

60

358

50

r

P

12/26/28

13.4

34.0

8.06

54

60

359

r

P

12/26/28

15.0

34.0

8.07

50

60-61

360

c

n

12/26/28

(15.0

34.0

8.07

50)

60-61

361

oc

n

12/26/28

115.0

34.0

8,07

50)

60-61

362

r

n

12/26/28

15.0

34.0

8.07

50)

61

363

r

n

12/28/28

16.9

34.0

8.05

46

61

364

50

r

n

2/28/28

14.0

34.0

8.05

. 60

61

365

100

r

n

2/28/28

10.8

34.0

8.03

80

61

366

a

P I

2/28/28

16.9

34.0

8.05

46

61

367

50

oc

P

2/28/28

14.0

34.0

8.05

60

61

368

100

oc

P 1

2/28/28

10.8

34.0

8.03

80

61-62

369

c

n

2/29/28

16.9
(16.9

34.0

8.05

46)
46

61-62

370

a

n

2/29/28

34.0

8.05

62

372

50

oc

n :

2/30/28

16.2

34.3

8.10

28

62

373

100

c

n ]

2/30/28

13.1

34.2

8.06

48

62

374

oc

P ]

2/30/28

19.2

34.2

8.12

32

62

375

50

r

P 1

2/30/28

16.2

34.3

8.10

28

62

376

100

r

P 1

2/30/28

13.1

34.2

8.06

48

62-63

377

r

n 1

2/31/28

(20.5

34.6

8.07

21)

63

382

r

P

1/ 1/29

20.5

34.6

8.07

21

63

384

100

r

P

1/ 1/29

15.6

34.6

8.08

24

63-64

386

00

n

1/ 1/29

20.5

34.6

8.07

21)

63-64

387

r

n

1/ 2/29

20.5

34.6

8.07

21

63-64

388

r

n

1/ 2/29

20.5

34.6

8.07

21

63-64

389

oc

n

1/ 2/29

20.5

34.6

8.07

21

63-64

390

r

n

1/ 2/29

20.5

34.6

8.07

21

63-64

391

oc

n

1/ 3/29

20.5

34.6

8.07

21

63-64

392

r

n

1/ 3/29

20.5

34.6

8.07

21

63-64

393

r

n

1/ 3/29

20.5

34.6

8.07

21

64

394

r

n

1/ 3/29

20.5

34.6

8.07

21)

64

395

50

r

n

1/ 3/29

17.2

34.6

8.12

29

64

397

100

r

n

1/ 3/29

15.8

34.5

8.10

32

64

398

oc

p

1/ 3/29

20.6

34.6

8.12

21

64

399

50

r

p

1/ 3/29

17.2

34.6

8.12

29

64

400

100

r

p

1/ 3/29

15.8

34.5

8.10

32

64-65

401

oc

n

1/ 3/29

20.6

34.6

8.12

21)

64-65

402

oc

n

1/ 4/29

20.6

34.6

8.12

21

64-65

403

oc

n

1/ 4/29

20.6

34.6

8.12

21

64-65

404

r

n

1/ 4/29 (

20.6

34.6

8.12

21

65

405

r

n

1/ 5/29

20.2

34.5

8.10

24

65

406

50

r

n

1/ 5/29

16.5

34.5

8.10

25

65

408

oc

p

1/ 5/29

20.2

34.5

8.10

24

65

409

50

oc

p

1/ 5/29

16.5

34.5

8.10

25

68

422

r

p

1/10/29

19.2

35.1

8.14

29

68

423

50

r

p

1/10/29

18.2

35.0

8.14

29

69

425

50

r

n

1/12/29

17.4

35.1

7.99

151

60

426

100

r

n

1/12/29

14.6

34.8

7.86

198

69

427

oc

P

1/12/29

21.1

35.2

8.12

62

70

432

50

r

n

1/13/29

15.4

35.0

7.88

178

70

433

100

r

n

1/13/29

12.6

34.8

7.68

233

70

434

r

P

1/13/29

21.2

35.1

8.05

103

58

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 6. Distributional and environmental records for Ceratlum furca Ehrenbergi- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/m3

71

439

50

r

n

2/ 6/29

16.7

35.1

7.90

150

71

441

00

p

2/ 6/29

23.5

35.2

8.13

58

72

444

50

oc

n

2/ 8/29

18.7

35.4

8.12

60

72

445

100

r

n

2/ 8/29

14.8

35.0

7.94

154

72

446

r

p

2/ 8/29

24.9

35.3

8.16

50

72

447

50

r

p

2/ 8/29

18.7

35.4

8.12

60

72-73

449

oc

n

2/ 9/29

(25.3

35.4

8.21

44)

73

450

oc

n

2/10/29

25.3

35.4

8.21

44

73

451

50

oc

n

2/10/29

18.7

35.4

8.05

122

73

452

100

oc

n

2/10/29

14.7

35.0

7.80

178

73

453

c

p

2/10/29

25.3

35.4

8.21

44

73

454

50

r

p

2/10/29

18.7

35.4

8.05

122

73

455

100

r

p

2/10/29

14.7

35.0

7.80

178

74

456

r

n

2/12/29

24.2

35.6

8.17

68

74

457

50

r

n

2/12/29

19.2

35.4

8.06

80

74

459

r

p

2/12/29

24.2

35.6

8.17

68

74

460

50

r

p

2/12/29

19.2

35.4

8.06

80

75

464

100

oc

n

2/14/29

17.8

35.4

8.00

75

75

465

r

p

2/14/29

22.8

35.8

8.18

44

77

473

r

p

2/18/29

23.7

36.0

8.19

16

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

478

r

P

2/20/29

24.6

36.0

8.17

32

78

479

50

r

p

2/20/29

23.8

36.1

8.14

32

79

483

100

r

n

2/22/29

21.8

36.2

8.13

45

80

488

r

p

2/24/29

26.0

35.9

8.20

36

80

490

100

r

p

2/24/29

23.4

36.2

8.16

32

81

493

r

p

2/26/29

26.5

35.8

8.19

38

82

498

r

p

2/28/29

27.2

36.3

8.21

34

83

504

50

r

p

3/ 2/29

27.4

36.5

8.24

25

89

530

r

P

3/23/29

28.4

35.6

8.25

21

89

532

r

n

3/23/29

(28.4

35.6

8.25

21)

Samoa

555

oc

n

4/ 2/29

96

568

r

n

4/26/29

29! 3

35! 3

8.23

12

95

568A

r

n

4/25/29

(29.4

34.7

8.26

14)

96

571

oc

p

4/26/29

29.3

35.3

8.23

12

96

573

100

oc

p

4/26/29

28.2

35.7

8.19

25

97

577

oc

p

4/28/29

28.3

35.2

8.16

24

97

578

50

r

p

4/28/29

28.0

35.4

8.16

21

97

579

100

oc

p

4/28/29

27.6

35.6

8.15

25

98

584

r

p

4/30/29

27.0

35.3

8.16

24

99

593

50

r

p

5/ 2/29

27.8

34.9

8.22

12

100

596

50

r

n

5/ 4/29

27.6

34.7

8.21

10

104

624

r

P

5/13/29

26.1

35.2

8.24

7

105

630

r

P

5/15/29

26.9

34.9

8.23

5

108

649

r

p

5/27/29

28.4

35.0

8.25

4

110

661

c

n

5/31/29

23.9

34.7

8.18

5

110

662

50

c

n

5/31/29

18.4

34.8

8.16

7

110

663

100

c

n

5/31/29

17.9

34.7

8.14

11

110

664

a

P

5/31/29

23.9

34.7

8.18

5

110

665

50

oc

P

5/31/29

18.4

34.8

8.16

7

110

666

100

oc

P

5/31/29

17.9

34.7

8.14

11

111

668

c

n

6/ 3/29

20.1

34.5

8.18

5

111

669

50

oc

n

6/ 3/29

19.4

34.6

8.17

5

111

671

oc

p

6/ 3/29

20.1

34.5

8.18

5

111

672

50

r

p

6/ 3/29

19.4

34.6

8.17

5

112

675

50

oc

n

6/ 5/29

21.7

34.6

8.23

7

112

676

100

oc

n

6/ 5/29

19.8

34.7

8.20

8

112

677

r

P

6/ 5/29

23.2

34.6

8.22

7

112

678

50

r

p

6/ 5/29

21.7

34.6

8.23

7

113

680

r

n

'6/25/29

24.2

34.5

8.25

5

113

683

r

p

6/25/29

24.2

34.5

8.25

5

114

688

100

r

P

6/27/29

13.0

34.5

8.00

91

114

689

c

p

6/27/29

19.9

34.3

8.15

7

114

690

50

oc

P

6/27/29

16.2

34.6

8.04

63

115

693

50

r

n

6/29/29

17.5

34.6

8.12

17

115

694

100

r

n

6/29/29

15.6

34.6

8.08

27

115

695

oc

p

6/29/29

20.6

34.6

8.19

4

115

696

oc

p

6^29/29

20.6

34.6

8.19

4

115

697

50

r

• P

6/29/29

17.5

34.6

8.12

17

117

709

50

r

n

7/ 3/29

12.5

34.2

8.06

51

117

710

100

r

n

7/ 3/29

8.8

34.1

7.98

84

117

711

r

P

7/ 3/29

15.9

34.3

8.17

3

129

780

r

n

7/27/29

16.3

33.1

8.13

25

130

784

r

p

9/ 4/29

16.2

33.4

8.34

36

131

792

r

P

9/ 4/29

16.2

33.4

8.34

36

APPENDIX

59

Table 6. Distributional and environinental records for Ceratium furca Ehrenbergi- -Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
("C)

Salinity
(o/oo) ,

pH

P04
mg/in3

131

793

r

P

9/ 6/29

19.3

33.4

8.34

131

794

r

P

9/ 6/29

19.3

33.4

8.34

>••

131

795

r

P

9/ 6/29

19.3

33.4

8.34

132

797

p

n

9/ 3/29

21.0

33.9

8.34

15

132

801
803

oc

P

S/ 8/29

21.0

33.9

8.34

15

132

100

r

p

9/ 8/29

14.3

33.4

8.30

16

136

832

r

P

9/16/29

24.6

35.4

8.37

3

136

833

50

r

p

9/16/29

21.4

35.1

8.39

3

140

858

100

r

n

10/ 3/29

25.5

35.0

8.34

7

141

869

50

r

P

10/ 5/29

24.8

35.3

8.34

5

142

872

p

n

10/ 7/29

24.1

34.8

8.33

5

142

877

50

r

P

10/ 7/29

21.8

34.8

8.30

5

142

879

100

p

P

10/ 7/29

16.6

34.4

8.27

7

144

889

100

r

P

10/11/29

16.6

34.5

8.37

6

146

897

50

r

n

10/15/29

22.4

34.9

8.30

6

146

900

. r

p

10/15/29

22.4

34.9

8.37

6

147

907

p

P

10/17/29

23.3

35.3

8.26

8

153

955

oc

P

10/29/29

28.1

34.2

8.47

7

158

983

r

n

11/ 8/29

28.2

35.6

8.34

36

158

985

100

r

n

11/ 8/29

27.6

35.9

8.39

48

Table 7. Distributional and environmental records for Ceratium belone

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

lb

8

100

r

n

5/16/28

20.3

36.5

8.18

36

18

114

p

p

8/17/28

27.0

37.0

8.23

5

47

267

50

p

n

11/23/28

23.8

36.0

8.23

20

47

268

100

p

n

11/23/28

22.7

36.2

8.23

20

49

277

oc

n

11/27/28

23.4

36.2

8.27

13

49

278

50

r

n

11/27/28

22.6

36.1

8.26

13

49

279

100

oc

n

11/27/28

21.6

35.9

8.26

13

49

280

p

p

11/27/28

23.4

36.2

8.27

13

49a

283

oc

n

11/28/28

(23.2

36.0

8.23

13)

50-51

290

r

n

11/30/28

22.8

35.6

8.22

16

50-51

291

oc

n

11/30/28

(22.8

35.6

8.22

16

50-51

292

r

n

11/30/28

(22.8

35.6

8.22

16

51

293

r

n

12/ 1/28

22.8

35.6

8.22

16

51

294

50

oc

n

12/ 1/28

20.5

35.6

8.22

17

51

295

100

oc

n

12/ 1/28

20.0

35.6

8.22

17

53-54

310

r

n

12/10/28

(23.0

35.6

8.22

11)

53-54

311

r

n

12/10/28

23.0

35.6

8.22

11)

53-54

313

r

n

12/10/28

(23.0

35.6

8.22

11)

54

320

50

r

n

12/14/28

19.8

35.4

8.18

17

54

321

100

p

n

12/14/28

18.7

35.4

8.16

20

55

323

p

n

12/16/28

20.4

34.9

8.19

12

55

324

50

p

n

12/16/28

18.7

35.0

8.18

12

56

330

100

p

n

12/18/28

16.6

34.8

8 11

12

74

456

p

n

2/12/29

24.2

35.6

8.17

68

78

481

p

n

2/22/29

25.2

36.0

8.17

34

79

482

50

p

n

2/22/29

24.5

36.1

8.17

34

79

483

100

p

n

2/22/29

21.8

36.2

8.13

45

80

488

oc

P

2/24/29

26.0

35.9

8.20

36

81

493

p

p

2/26/29

26.5

35.8

8.19

38

82

496

p

n

2/28/29

27.2

36.3

8.21

34

91

540

p

n

3/27/29

28.7

35.1

8.30

21

91

541

50

p

n

3/27/29

28.5

35.2

8.30

24

94

558

50

p

n

4/22/29

29.3

34.7

8.25

14

95

563

50

p

n

4/24/29

29.3

34.9

8.24

16

96

569

50

r

n

4/26/29

29.2

35.3

8.23

25

96

570

100

■ p

n

4/26/29

28.2

35.7

8.19

25

96

571

p

P

4/26/29

29.3

35.3

8.23

12

97

574

p

n

4/28/29

28.3

35.2

8.16

24

97

576

100

p

n

4/28/29

27.6

35.6

8.15

25

97

577

p

P

4/28/29

28.3

35.2

8.16

24

97

579

100

p

P

4/28/29

27.6

35.6

8.15

25

100

598

p

p

5/ 4/29

27.7

34.7

8.21

10

133

804

p

n

9/10/29

22.7

34.7

8.47

7

142

874

100

p

n

10/ 7/29

16.6

34.4

8.27

7

145

890

p

n

10/13/29

22.3

34.6

8.29

6

145

893

r

p

10/13/29

22.3

34.6

8.29

6

145

895

100

r

P

10/13/29

16.0

34.1

3.31

6

60

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 7. Distributional and environmental records for Ceratium belone--Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

PO4

mg/m3

158

983

r

n

11/ 8/29

28.2

35.6

8.34

36

158

985

100

r

n

11/ 8/29

27.6

35.9

8.39

48

159

990

r

n

11/11/29

28.6

35.7

8.37

15

159

991

50

r

n

11/11/29

28.5

35.7

8.39

15

Table 8. Distributional and environmental records for Ceratium incisum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(0/00)

pH

PO4

mg/m^

16

103

r

P

8/13/28

25.9

36.2

8.24

8

33

198

50

r

n

10/ 8/28

28.2

36.2

8.24

4

33

198A

50

r

p

10/ 8/28

28.2

36.2

8.24

4

40

232

r

p

11/ 8/28

22.2

33.7

8.21

24

46

261

r

n

11/21/28

23.3

35.3

8.16

36

46

261

r

p

11/21/28

23.3

35.3

8.16

36

78

477

100

r

n

2/20/29

21.9

36.2

8.14

34

79

483

100

r

n

2/22/29

21.8

36.2

8.13

45

80

488

r

P

2/24/29

26.0

35.9

8.20

36

81

493

r

p

2/26/29

26.5

35.8

8.19

38

82

497

50

r

n

2/28/29

27.2

36.3

8.21

34

94

559

100

r

n

4/22/29

28.5

35.6

8.21

25

95

563

50

r

n

4/24/29

29.3

34.9

8.24

16

96

570

100

r

n

4/26/29

28.2

35.7

8.19

25

97

574

r

n

4/28/29

28.3

35.2

8.16

24

98

588

0-100

r

p

4/30/29

26.7

35.4

8.14

32

99

591

100

r

n

5/ 2/29

27.8

35.0

8.22

17

100

597

100

r

n

5/ 4/29

27.6

34.7

8.22

12

100

602

100

r

P

5/ 4/29

27.6

34.7

8.22

12

101

605

100

r

n

5/ 7/29

25.2

35.1

8.23

8

103

615

r

n

5/11/29

26.0

35.0

8.25

5

105

628

50

r

n

5/15/29

26.8

34.9

8.23

5

106

634

50

r

n

5/17/29

27.0

35.0

8.23

5

108

648

100

r

n

5/27/29

25.2

35.0

8.23

4

139

850

50

r

n

9/22/29

25.8

34.9

8.31

6

141

864

r

n

10/ 5/29

25.9

35.2

8.34

5

152

945

50

r

n

10/27/29

14.2

34.5

7.87

53

155

965

r

n

11/ 2/29

27.8

34.9

8.29

29

156

973

50

r

n

11/ 4/29

27.0

35.1

8.37

46

158

985

100

r

n

11/ 8/29

27.6

35.9

8.39

48

Table 9. Distributional and environmental records for Ceratium pentagonum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(0/00)

pH

PO4
mg/m3

Subspe

cies tene

rum

13a

88

c

n

8/ 8/28

(21.2
(21.2

35.2

8.18

11

13a

89

r

n

8/ 8/28

35.2

8.18

11

14

90

r

n

8/ 9/28

21.2

35.2

8.18

11

14

92

50

r

P

8/ 9/28

15.0

35.1

8.18

16

14

93

oc

P

8/ 9/28

21.2

35.2

8.18

U

15

97

r

P

8/11/28

24.8

36.4

8.21

11

15

98

50

r

P

8/11/28

19.8

36.5

8.21

8

15

99

100

r

p

8/11/28

18.9

36.4

8.20

19

16

103

r

P

8/13/28

25.9

36.2

8.24

8

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

17

109

50

oc

P

8/15/28

21.9

36.6

8.28

12

17

110

50

oc

P

8/15/28

21.9

36.6

8.28

12

17

111

oc

n

8/15/28

26.2

36.6

8.29

9

18

114

r

P

8/17/28

27.0

37.0

8.23

5

18

115

50

oc

p

8/17/28

22.4

36.8

8.24

5

18

118

50

r

n

8/17/28

22.4

36.8

8.24

5

19

121

50

r

P

8/20/28

25.2

37.1

8.27

5

20

125

r

p

8/22/28

26.0

36.6

8.37

5

20

126

50

r

P

8/22/28

25.8

36.6

8.26

3

20

127

100

r

p

8/22/28

22.6

36.7

8.19

5

20

128

r

n

8/22/28

26.0

36.6

8.37

5

APPENDIX

61

Table 9. Distributional and environmental records for Ceratiuin pentagonum- -Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

PO4
mg/m3

20

130

100

oc

n

8/22/28

21

134

100

r

P

8/24/28

21

136

50

r

n

8/24/28

22

141

100

r

P

8/27/28

23

145

r

P

8/29/28

24

155

50

r

n

8/31/28

25

157

r

P

9/ 3/28

26

164

50

r

P

9/ 5/28

30

189

50

r

n

9/15/28

30

190

100

r

n

9/15/28

37

221

50

r

P

11/ 1/28

40

231

100

r

n

11/ 8/28

40

232

oc

P

11/ 8/28

40

234

100

r

P

11/ 8/28

41

235

r

n

11/10/28

41

236

50

oc

n

11/10/28

41

238

r

P

11/10/28

41

239

100

r

P

11/10/28

42

241

r

n

11/13/28

42

242

50

r

n

11/13/28

43

245

50

r

n

11/15/28

43

246

100

r

n

11/15/28

43

247

r

P

11/15/28

43

248

50

r

P

11/15/28

43

249

100

r

P

11/15/28

44

252

100

r

n

11/17/28

44

253

oc

P

11/17/28

44

254

50

oc

P

11/17/28

44

255

100

r

P

11/17/28

45

256

r

n

11/19/28

45

257

50

oc

n

11/19/28

45

258

100

r

n

11/19/28

45

259

oc

P

11/19/28

45

260

50

r

P

11/19/28

46

264

oc

P

11/21/28

46

265

50

r

P

11/21/28

47

269

r

P

11/23/28

47

270

50

r

P

11/23/28

48

274

r

p

11/23/28

49

280

r

P

11/27/28

49

281

50

r

P

11/27/28

50

286

100

r

n

11/29/28

50

287

r

P

11/29/28

50

288

50

r

P

11/29/28

50

289

100

r

P

11/29/28

54

322

r

P

11/29/28

59

348

r

n

12/24/28

60

359

r

p

12/26/28

62

371

r

n

12/30/28

62

372

50

r

n

12/30/28

62

373

100

oc

n

12/30/28

62

376

100

r

P

12/30/28

63

377

r

n

12/31/28

63

380

50

r

n

1/ 1/29

63

381

100

r

n

1/ 1/29

63

383

50

r

P

1/ 1/29

63-64

385

r

n

1/ 1/29

63-64

386

oc

n

1/ 1/29

63-64

387

r

n

1/ 2/29

63-64

389

r

n

1/ 2/29

63-64

390

r

n

1/ 2/29

63-64

391

oc

n

1/ 3/29

63-64

392

oc

n

1/ 3/29

64

397

100

r

n

1/ 3/29

64

398

r

P

1/ 3/29

64

400

100

r

P

1/ 3/29

64-65

401

r

n

1/ 3/29

64-65

402

r

n

1/ 4/29

65

409

50

r

P

1/ 5/29

66

411

r

n

1/ 7/29

66

412

100

r

n

1/ 7/29

66

413

oc

P

1/ 7/29

66

414

50

r

P

1/ 7/29

67

416

r

n

1/ 8/29

Subspecies tenerum- -Continued

22.6
21.0
24.4
17.5
27.2
23.1
27.5
24.1
27.8
24.1
18.8
13.9
22.2
13.9
20.4
14.6
20.4
14.5
18.7
17.2
17.0
13.6
19.6
17.0
13.6
13.8
20.7
20.4
13.8
22.4
22.4
18.6
22.4
22.4
23.3
23.2
23.9
23.8
23.6
23.4
22.6
20.5
23.2
22.0
20.5
23.4
16.3
15.0
19.2
16.2
13.1
13.1
(20.5
17.0
15.6
U.O

i20.5
20.5
20.5
20.5
20.5
20.5
20.5
15.8
20.6
15.8
(20.6
(20.6
16.5
19.4
17.8
19.4
17.8
19.3

36.7

8.19

5

36.8

8.20

7

36.2

8.26

4

36.1

7.99

123

35.9

8.25

4

36.0

8.14

8

35.6

8.31

5

36.1

8.21

5

36.1

8.29

5

36.4

8.10

20

34.5

8.00

121

35.0

7.85

159

33.7

8.21

24

35.0

7.85

159

34.2

8.11

32

35.0

7.94

58

34.2

8.11

32

35.0

7.91

152

34.7

8.06

45

34.9

7.99

68

34.9

7.93

80

35.0

7.90

92

34.8

8.09

52

34.9

7.93

80

35.0

7.90

92

35.0

7.85

70

34.9

8.03

38

34.9

8.04

34

35.0

7.85

70

35.3

8.12

38

35.2

8.13

46

35.1

8.00

50

35.3

8.12

38

35.2

8.13

46

35.3

8.16

36

35.3

8.16

40

36.0

8.16

17

36.0

8.16

20

36.4

8.23

13

36.2

8.27

13

36.1

8.26

13

35.7

8.22

13

36.0

8.23

13

35.9

8.23

13

35.7

8.22

13

35.5

8.22

9

34.0

8.10

38

34.0

8.07

50

34.2

8.12

32

34.3

8.10

28

34.2

8.06

48

34.2

8.06

48

34.6

8.07

21)

34.6

8.08

25

34.6

8.08

24

34.6

8.08

25

34.6

8.07

21)

34.6

8.07

21

34.6

8.07

21)

34.6

8.07

21

34.6

8.07

21

34.6

8.07

21

34.6

8.07

21)

34.5

8.10

32

34.6

8.12

21

34.5

8.10

32

34.6

8.12

21)
21)

34.6

8.12

34.5

8.10

25

34.7

8.10

29

34.9

8.12

21

34.7

8.10

29

34.8

8.10

29

34.9

8.11

21

62

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 9. Distributional and environmental records for Ceratium pentagonum- -Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

PO4
mg/m3

Subspecies tenerum- -Continued

67

418

100

r

n

1/ 8/29

16.2

34.6

8.05

40

68

422

r

p

1/10/29

19.2

35.1

8.14

29

68

423

50

r

P

1/10/29

18.2

35.0

8.14

29

69

425

50

r

n

1/12/29

17.4

35.1

7.99

151

69

426

100

r

n

1/12/29

14.6

34.8

7.86

198

69

427

oc

p

1/12/29

21.1

35.2

8.12

62

69

428

50

r

P

1/12/29

17.4

35.1

7.99

151

70

432

50

r

n

1/13/29

15.4

35.0

7.88

178

70

434

r

P

1/13/29

21.2

35.1

8.05

103

70

436

100

r

p

1/13/29

12.6

34.8

7.68

233

71

441

oc

p

2/ 6/29

23.5

35.2

8.13

58

72

444

50

r

n

2/ 8/29

18.7

35.4

8.12

60

72

446

r

p

2/ 8/29

24.9

35.3

8.16

50

72

447

50

oc

P

2/ 8/29

18.7

35.4

8.12

60

72

448

100

r

p

2/ 8/29

14.8

35.0

7.94

154

73

450

r

n

2/10/29

25.3

35.4

8.21

44

73

451

50

oc

n

2/10/29

18.7

35.4

8.05

122

78

452

100

oc

n

2/10/29

14.7

35.0

7.80

178

73

453

oc

p

2/10/29

25.3

35.4

8.21

44

73

454

50

r

p

2/10/29

18.7

35.4

8.05

122

74

459

r

P

2/12/29

24.2

35.6

8.17

68

79

481

r

n

2/22/29

25.2

36.0

8.17

68

79

482

50

r

n

2/22/29

24.5

36.1

8.17

68

79

485

50

oc

P

2/22/29

24.5

36.1

8.17

68

87

522

50

r

n

3/11/29

26.5

36.1

8.26

20

87

523

r

P

3/11/29

27.8

36.1

8.28

17

87

524

50

r

P

3/11/29

26.5

36.1

8.26

20

87

525

100

r

P

3/11/29

23.9

36.0

8.23

20

89

531

100

r

p

3/23/29

26.4

36.0

8.24

12

90

533

r

n

3/25/29

28.5

35.5

8.27

21

90

535

r

p

3/25/29

28.5

35.5

8.27

21

90

536

50

r

P

3/25/29

28.6

35.6

8.26

21

91

541

50

r

n

3/27/29

28.5

35.2

8.30

24

91

542

r

p

3/27/29

28.7

35.1

8.30

21

91

543

50

r

P

3/27/29

28.5

35.2

8.30

24

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

553

100

r

p

3/31/29

27.6

35.8

8.27

29

96

572

50

r

P

4/26/29

29.2

35.3

8.23

12

101

606

r

p

5/ 7/29

26.3

35.3

8.24

8

102

612

r

P

5/ 9/29

25.8

35.0

8.24

8

102

613

50

r

p

5/ 9/29

25.8

35.0

8.24

8

104

624

oc

P

5/13/29

26.1

35.2

8.24

7

104

626

100

oc

P

5/13/29

25.3

35.3

8.21

7

105

630

oc

P

5/19/29

26.9

34.9

8.23

5

105

631

50

r

P

5/19/29

26.8

34.9

8.23

5

106

635

100

r

n

5/17/29

25.6

35.1

8.23

5

106

636

r

P

5/17/29

27.2

35.0

8.23

5

106

637

50

r

P

5/17/29

27.0

35.0

8.23

5

107

642

r

P

5/19/29

28.0

34.4

8.23

5

107

645

50

oc

p

5/19/29

27.9

34.4

8.23

4

108

647

50

r

n

5/27/29

26.8

35.0

8.24

4

108

648

100

r

n

5/27/29

25.2

35.0

8.23

4

110

665

50

oc

P

5/31/29

18.4

35.0

8.16

7

110

666

100

oc

P

5/31/29

17.9

34.7

8.14

11

111

671

oc

P

5/31/29

17.9

34.7

8.14

11

111

672

50

r

P

5/31/29

19.4

34.6

8.17

11

112

675

50

r

n

6/ 5/29

21.7

34.6

8.23

7

112

677

r

p

6/ 5/29

23.2

34.6

8.22

7

112

678

50

r

P

6/ 5/29

21.7

34.6

8.23

7

Subspecies pacificum

118
120

719
729

"0

r
r

P

7/ 5/29
7/ 9/29

"7!2

33!b

7.98

137

120

730

50

r

p

7/ 9/29

2.2

33.1

7.90

177

121

732

r

n

7/11/29

7.5

32.9

7.98

141

121

735

oc

p

7/11/29

7.5

32.9

7.98

141

122

738

r

p

7/13/29

8.2

32.8

7.98

130

122

741

r

n

7/13/29

8.2

32.8

7.98

130

123

745

50

oc

P

7/15/29

4.4

32.9

7.94

150

123

747

r

n

7/15/29

8.1

32.8

8.03

113

124

749

oc

n

7/17/29

9.3

32.7

8.04

113

124

752

r

p

7/17/29

9.3

32.7

8.04

103

124

750

50

r

n

7/17/29

5.4

32.7

8.02

110

12s

755

c

n

7/19/29

10.5

32.8

8.03

125

APPENDIX

63

Table 9. Distributional and environmental records for Ceratium pentagonum- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

Subspecies pa(

:if icum - -Concluded

125

756

50

DC

n

7/19/29

5.5

32.8

7.98

138

125

757

100

oc

n

7/19/29

4.2

32.9

7.90

175

125

758

c

P

7/19/29

10.5

32.8

8.03

125

125

759

50

oc

P

7/19/29

5.5

32.8

7.98

138

Subspecies tenerum- -Concluded

132

803

100

r

P

9/ 8/29

14.3

33.4

8.30

16

133

804

r

n

9/10/29

22.7

34.7

8.47

7

134

815

r

P

9/12/29

22.9

34.7

8.34

6

135

821

50

r

n

9/14/29

21.5

35.0

8.37

5

135

825

50

r

P

9/14/29

21.5

35.0

8.37

5

135

826

100

r

P

9/14/29

18.7

34.8

8.34

5

136

829

100

r

n

9/16/29

18.6

35.0

8.39

3

136

833

50

r

P

9/16/29

21.4

35.1

8.39

3

138

847

r

P

9/20/29

26.1

34.8

8.35

3

138

848

50

r

P

9/20/29

25.6

34.7

8.30

3

139

850

50

r

n

9/22/29

25.8

34.9

8.31

6

139

851

100

r

n

9/22/29

22.4

35.2

8.28

6

139

854

50

r

p

9/22/29

25.8

34.9

8.31

6

139

855

100

r

p

9/22/29

22.4

35.2

8.28

6

141

864

r

n

10/ 5/29

25.9

35.2

8.28

5

141

865

50

r

n

10/ 5/29

24.8

35.3

8.28

5

141

871

100

r

P

10/ 5/29

20.0

35.0

8.33

5

143

884

r

p

10/ 9/29

22.4

34.4

8.30

6

143

885

50

r

p

10/ 9/29

19.0

34.2

8.34

6

146

900

r

P

10/15/29

22.4

34.9

8.37

6

146

902

100

r

P

10/15/29

19.7

34.3

8.26

7

147

904

50

r

n

10/17/29

23.1

35.3

8.29

5

147

907

r

P

10/17/29

23.3

35.3

8.26

8

147

908

50

r

P

10/17/29

19.2

35.0

8.29

5

148

911

50

r

n

10/19/29

23.0

35.1

148

917

oc

P

10/19/29

23.4

35.2

• • •

149

921

50

r

n

10/21/29

23.3

35.0

8.37

6

149

927

oc

p

10/21/29

23.5

35.0

8.34

6

150

931

100

r

n

10/23/29

19.6

34.6

8.32

6

150

936

100

r

P

10/23/29

19.6

34.6

8.32

6

151

941

r

p

10/26/29

26.0

34.0

152

946

100

r

n

10/27/29

11.4

34.7

7.76

75

152

948

r

P

10/27/29

27.4

33.7

8.35

20

Table 10. Distributional and environmental records for Ceratium subrobustum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(0/00)

pH

PO4
mg/m3

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

19

123

50

oc

n

8/20/28

25.2

27.1

8.27

5

47

268

100

r

n

11/23/28

22.7

36.2

8.23

20

48

273

100

oc

n

11/25/28

22.7

36.3

8.26

16

48

276

100

r

p

11/25/28

22.7

36.3

8.26

16

49

279

100

oc

n

n/27/28

21.6

35.9

8.26

13

49

282

100

r

p

11/27/28

21.6

35.9

8.26

13

50

286

100

r

n

11/29/28

20.5

35.7

8.22

13

51

298

100

r

p

12/ 1/28

20.0

35.6

8.22

17

53

305

100

r

n

12/ 5/28

19.9

35.6

8.19

13

54

321

100

r

n

12/14/28

18.7

35.4

8.16

20

55

324

50

oc

n

12/16/28

18.7

35.0

8.18

12

55

325

100

r

n

12/16/28

16.7

34.9

7.17

12

56

330

100

oc

n

12/18/28

16,6

34.8

8.11

12

57

337

100

oc

n

12/20/28

14.3

34.4

8.10

40

57

340

100

r

P

12/20/28

14.3

34.4

8.10

40

58

342

50

r

n

12/22/28

14.8

34.0

8.12

25

58

345

50

r

p

12/22/28

14.8

34.0

8.12

25

75

462

r

n

2/14/29

22.8

35.8

8.18

44

76

468

50

r

n

2/16/29

22.1

35.9

8.14

42

76

469

100

r

n

2/16/29

21.2

35.8

8.12

45

77

472

r

n

2/18/29

23.7

36.0

8.19

16

77

473

r

p

2/18/29

23.7

36.0

8.19

16

77

474

50

r

p

2/18/29

23.5

36.0

8.19

16

78

477

100

r

n

2/20/29

24.6

36.0

8.17

32

94

559

100

r

n

4/22/29

28.5

35.6

8.21

25

64

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 10. Distributional and environmental records for Ceratium subrobustxun- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

Cc)

(o/oo)

mg/m3

97

575

50

r

n

4/28/29

28.0

35.4

8.16

21

97

576

100

r

n

4/28/29

27.6

35.6

8.15

25

97

577

r

p

4/28/29

28.3

35.2

8.16

24

97

579

100

r

P

4/28/29

27.6

35.6

8.15

25

98

584

r

P

4/30/29

27.0

35.3

8.16

24

98

585

50

r

p

4/30/29

26.9

35.3

8.16

28

99

589

r

n

5/ 2/29

27.9

34.9

8.21

12

99

590

50

r

n

5/ 2/29

27.8

34.9

8.22

12

99

591

100

r

n

5/ 2/29

27.8

35.0

8.22

17

99

592

oc

P

5/ 2/29

27.9

34.9

8.21

12

99

595

r

n

5/ 2/29

27.9

34.9

8.21

12

100

596

50

r

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

oc

n

5/ 4/29

27.6

34.7

8.22

12

100

599

50

r

P

5/ 4/29

27.6

34.7

8.21

10

100

602

100

r

P

5/ 4/29

27.6

34.7

8.22

12

109

656

100

r

n

5/29/29

19.4

34.8

8.18

5

109

658

50

r

P

5/29/29

23.1

35.0

8.22

3

109

659

100

r

P

5/29/29

19.4

34.8

8.18

5

111

668

r

n

6/ 3/29

20.1

34.5

8.18

5

113

684

50

r

P

6/25/29

23.8

34.6

8.25

5

114

689

r

P

6/27/29

19.9

34.3

8.15

7

115

695

r

P

6/29/29

20.6

34.6

8.19

4

133

805

50

r

n

9/10/29

20.8

34.7

8.37

7

134

817

100

r

p

9/12/29

18.1

34.6

8.34

6

136

829

100

r

n

9/16/29

18.6

35.0

8.39

3

137

841

50

r

P

9/18/29

29.4

35.1

8.34

4

139

851

100

r

n

9/22/29

22.4

35.2

8.28

6

139

855

100

r

P

9/22/29

22.4

35.2

8.28

6

140

858

100

r

n ]

0/ 3/29

25.5

35.0

8.34

7

140

862

100

r

P ]

10/ 3/29

25.5

35.0

8.34

7

142

873

50

r

n ]

10/ 7/29

21.8

34.8

8.30

5

142

874

100

c

n 10/ 7/29

16.6

34.4

8.27

7

142

878

100

r

p 10/ 7/29

16.6

34.4

8.27

7

142

879

100

r

p 10/ 7/29

16.6

34.4

8.27

7

144

888

50

oc

p 10/11/29

21.1

34.7

8.33

6

144

889

100

oc

p 10/11/29

16.6

34.5

8.37

6

145

891

50

c

n 10/13/29

18.7

34.3

8.34

6

145

892

100

oc

n 10/13/29

16.0

34.1

8.31

6

145

894

50

r

p 10/13/29

18.7

34.3

8.34

6

145

895

100

r

p 10/13/29

16.0

34.1

8.31

6

146

897

50

oc

n 10/15/29

22.4

34.9

8.30

6

147

905

100

r

n 10/15/29

19.7

34.3

8.26

7

149

922

100

r

n 10/15/29

19.7

34.3

8.26

7

151

939

100

r

n 10/26/29

12.5

34.6

153

957

100

r

p 10/29/29

20.5

34.7

8.28

31

155

965

r

n 11/ 2/29

27.8

34.9

8.29

29

155

966

50

r

n 11/ 2/29

27.7

34.9

8.30

30

155

967

100

oc

n 11/ 2/29

27.2

35.0

8.30

35

155

973

50

oc

p 11/ 2/29

27.7

34.9

8.30

30

156

972

r

n 11/ 4/29

27.6

35.0

8.34

28

156

974

100

r

n 11/ 4/29

26.4

35.1

8.30

48

157

978

c

n 11/ 6/29

27.1

35.3

8.27

47

157

979

50

oc

n 11/ 6/29

27.1

35.2

8.32

60

157

980

100

r

n 11/ 6/29

26.8

35.5

8.30

64

158

985

100

r

n 11/ 8/29

27.6

35.9

8.39

48

Table 11. Distributional and environmental records for Ceratium teres

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature

CC)

Salinity
(0/00)

pH

PO4
mg/m^

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

17

109

r

P

8/15/28

26.2

36.6

8.29

9

18

114

oc

P

8/17/28

27.0

37.0

8.23

5

18

115

50

r

P

8/17/28

22.4

36.8

8.24

5

18

118

50

r

n

8/17/28

22.4

36.8

8.24

5

19

121

50

r

P

8/20/28

25.2

37.1

8.27

5

19

123

50

r

n

8/20/28

25.2

37.1

8.27

5

20

127

100

r

P

8/22/28

22.6

36.7

8.19

5

20

130

100

r

n

8/22/28

22.6

36.7

8.19

5

20-21

131

r

n

8/23/28

(26.6

36.3

8.32

4)

21

134

100

r

P

8/24/28

21.0

36.8

8.20

7

APPENDIX

65

Table 11. Distributional and environmental records for Ceratlum teres- -Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

21

137

100

r

22

140

50

r

23

143

50

r

22

144

100

r

23

145

oc

23

149

50

r

24

151

r

24

152

50

r

30

189

50

r

32

194A

r

46

262

50

r

46

263

100

oc

46

264

r

47

270

50

r

48

276

100

r

49

279

100

r

49

281

50

r

50

289

100

oc

50-51

290

r

50-51

291

r

50-51

292

r

51

294

50

r

51

297

50

r

54

322

r

57

338

r

58

342

50

r

58

344

r

58

345

50

oc

59

348

r

59

350

100

oc

59

351

oc

59

352

SO

r

60

357

r

60-61

360

r

60-61

361

r

67

416

r

68

422

r

80.

489

50

r

80

490

100

r

81

493

r

81

495

100

r

85

512

r

87

523

r

87

524

50

r

89

530

r

90

536

50

r

90

537

100

r

91

542

oc

91

544

100

r

92

547

r

92

549

100

oc

93

552

oc

93

553

100

oc

94

562

100

r

95

565

r

95

566

50

r

95

567

100

r

96

573

100

r

97

577

oc

97

578

50

oc

99

593

50

r

100

598

r

100

602

100

oc

101

606

r

102

610

50

r

102

612

r

102

614

100

oc

103

616

50

r

103

618

oc

103

620

100

r

104

626

100

oc

105

630

oc

105

631

50

oc

107

640

50

r

107

642

oc

n

8/24/28

21.0

P

8/27/28

24.5

n

8/27/28

24.5

n

8/27/28

17.5

P

8/29/28

27.2

n

8/29/28

20.9

P

8/31/28

27.2

P

8/31/28

23.1

n

9/15/28

27.8

P

10/ 5/28

28.0

n

11/21/28

23.2

n

11/21/28

22.5

P

11/21/28

23.3

P

11/23/28

23.8

P

11/25/28

22.7

n

11/27/28

21.6

P

11/27/28

22.6

P

11/29/28

20.5

n

11/30/28

(22.8

n

11/30/28

(22.8

n

11/30/28

(22.8

n

12/ 1/28

20.5

P

12/ 1/28

20.5

P

12/14/28

23.4

P

12/20/28

19.0

n

12/22/28

14.8

P

12/22/28

17.0

P

12/22/28

14.8

n

12/24/28

16.3

n

12/24/28

11.4

P

12/24/28

16.3

P

12/24/28

14.0

P

12/26/28

15.0

n

12/26/28

(15.0
(15.0

n

12/26/28

n

1/ 8/29

19.3

P

1/10/29

19.2

P

2/24/29

25.9

P

2/24/29

23.4

P

2/26/29

26.5

P

2/26/29

23.6

P

3/ 6/29

27.9

P

3/11/29

27.8

P

3/11/29

26.5

P

3/23/29

28.4

P

3/25/29

28.6

P

3/25/29

26.3

P

3/27/29

28.7

P

3/27/29

25.8

P

3/29/29

28.5

P

3/29/29

26.2

P

3/31/29

28.7

P

3/31/29

27.6

P

4/22/29

28.5

P

4/24/29

29.4

P

4/24/29

29.3

P

4/24/29

28.5

P

4/26/29

28.2

P

4/28/29

28.3

P

4/28/29

28.0

P

5/ 2/29

27.8

P

5/ 4/29

27.7

P

5/ 4/29

27.6

P

5/ 7/29

26.3

n

5/ 9/29

25.8

P

5/ 9/29

25.8

P

5/ 9/29

25.6

n

5/11/29

25.8

P

5/11/29

26.0

P

5/11/29

24.8

P

5/13/29

25.3

P

5/15/29

26.9

P

5/15/29

26.8

n

5/19/29

27.9

P

5/19/29

28.0

36.8

36.2

36.2

36.1

35.9

36.0

35.2

36.0

36.1

36.0

35.3

35.4

35.3

36.0

36.3

35.9

36.1

35.7

35.6

35.6

35.6

35.6

35.6

35.5

34.5

34.0

34.0

34.0

34.0

34.1

34.0

34.0

34.0

34.0

34.0

34.9

35.1

36.0

36.2

35.8

36.2

36.2

36.1

36.1

35.6

35.6

35.8

35.1

36.0

35.3

36.0

34.7

35.8

35.6

34.7

34.9

35.4

35.7

35.2

35.4

34.9

34.7

34.7

34.7

35.0

35.0

35.0

35.2

35.0

35.2

35.3

34.9

34.9

34.4

34.4

8.20

8.21

8.21

7.99

8.25

8.14

8.32

8.14

8.29

8.23

8.16

8.17

8.16

8.23

8.26

8.26

8.26

8.22

8.22

8.22

8.22

8.22

8.22

8.22

8.14

8.12

8.12

8.12

8.10

8.03

8.10

8.08

8.07

8.07

8.07

8.11

8.14

8.19

8.16

8.19

8.18

8.22

8.28

8.26

8.25

8.26

8.24

8.30

8.25

8.29

8.28

8.30

8.27

8.21

8.26

8.24

8.22

8.19

8.16

8.16

8.22

8.21

8.22

8.24

8.24

8.24

8.23

8.25

8.25

8.25

8.21

8.23

8.23

8.23

8.23

7

9

9

123

4

13

4

8

3

2

40

40

36

20

16

13

13

13

16

16;

16
17
17
9
20
25
20
25
38
72
38
38
50
501
50)
21
29
29
32
38
36
40
17

2D

21

21

21

21

30

28

28

28

29

25

14

16

21

25

24

21

12

10

12
8
8
8
8
5
5
5
7
5
5
4
5

66

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 11. Distributional and environmental records for Ceratium teres — Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

109

660

r

p

5/29/29

27.4

35.0

8.23

3

112

675

SO

r

n

6/ 5/29

21.7

34.6

8.23

7

112

676

100

r

n

6/ 5/29

19.8

34.7

8.20

8

112

677

r

p

6/ 5/29

23.2

34.6

8.22

7

115

692

r

n

6/29/29

20.6

34.6

8.19

4

132

801

r

p

9/ 8/29

21.0

33.9

8.34

15

134

815

r

p

9/12/29

22.9

34.7

8.34

6

136

828

50

r

p

9/16/29

21.4

35.1

8.39

3

136

832

r

p

9/16/29

24.6

35.4

8.37

3

136

833

50

r

n

9/16/29

21.4

35.1

8.39

3

141

869

50

r

p

10/ 5/29

24.8

35.3

8.34

5

142

876

r

p

10/ 7/29

24.1

34.8

8.33

5

144

887

oc

p

10/11/29

23.3

35.0

8.37

6

145

893

r

p

10/13/29

22.3

34.6

8.29

6

146

901

50

r

p

10/15/29

22.4

34.9

8.30

6

146

902

100

r

p

10/15/29

19.7

34.3

8.26

7

147

907

r

p

10/17/29

23.3

35.3

8.26

8

148

918

50

r

p

10/19/29

23.0

35.1

151

941

r

p

10/26/29

26.0

34.0

153

956

50

r

P

10/29/29

28.1

34.4

8.39

7

Table 12. Distributional and environmental records for Ceratium kofoldii

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature

Cc)

Salinity
(o/oo)

pH

P04
mg/m3

44

253

45

256

45

257

50

45

259

71

441

100

600

100

111

671

111

672

50

112

679

100

130

784

130

785

50

142

879

100

r

r

r

oc

r

r

oc

r

r

r

r

r

11/17/28

20.7

34.9

8.03

38

11/19/28

22.4

35.3

8.12

38

11/19/28

22.4

35.2

8.13

46

11/19/28

22.4

35.3

8.12

38

2/ 6/29

23.5

35.2

8.13

58

5/ 4/29

27.6

34.7

8.22

12

6/ 3/29

20.1

34.5

8.18

5

6/ 3/29

19.4

34.6

8.17

5

6/ 5/29

19.8

34.7

8.20

8

9/ 4/29

16.2

33.4

8.34

36

9/ 4/29

11.7

33.4

8.26

83

10/ 7/29

16.6

34.4

8.27

7

Table 13. Distributional and environmental records for Ceratium bohmii

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4

mg/m3

40

230

50

'r

n

11/ 8/28

15.3

34.9

7.87

161

40

232

r

p

11/ 8/28

22.2

33.7

8.21

24

40

234

100

r

P

11/ 8/28

13.9

35.0

7.85

159

42

243

100

r

n

11/13/28

13.8

35.0

7.91

150

98

585

50

r

p

4/30/29

26.9

35.3

8.16

28

99

589

r

n

5/ 2/29

27.9

34.9

8.21

12

99

594

100

r

p

5/ 2/29

27.8

35.0

8.22

17

114

689

oc

P

6/27/29

19.9

34.3

8.15

7

114

690

50

r

P

6/27/29

16.2

34.6

8.04

63

115

695

oc

P

6/29/29

20.6

34.6

8.19

4

115

696

oc

p

6/29/29

20.6

34.6

8.19

4

115

698

100

oc

P

6/29/29

15.6

34.6

8.08

27

APPENDIX

67

Table 14. Distributional and environmental records for Ceratium lineatum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

PH

P04

mg/m3

3

20

r

n

5/21/28

15.5

36.1

8.15

99

4

23

100

r

n

5/23/28

13.4

35.9

8.12

50

6

37

100

r

n

5/31/28

11.3

35.6

8.08

41

6b

46

r

n

5/31/28

(12.4

35.6

8.15

21)

9

64

r

n

7/28/28

11.2

35.1

8.08

20

9

65

50

oc

n

7/28/28

8.4

35.1

7.96

55

9

66

100

r

n

7/28/28

7.6

35.1

7.98

56

10

68

oc

n

7/30/28

10.9

34.9

8.08

28

10

69

50

oc

n

7/30/28

10.0

34.9

8.04

34

10

70

100

oc

n

7/30/28

6.6

35.0

7.95

52

11a

78

.

oc

n

8/ 1/28

10.7

34.9

8.06

27

13a

88

r

n

8/ 9/28

21.2

35.2

8.18

11)

17

111

oc

n

8/16/28

26.2

36.6

8.29

9

19

122

r

n

8/20/28

26.6

37.0

8.34

5

115a

700

c

n

7/ 1/29

(18.4

34.3

8.18

4)

115a

702

oc

n

7/ 1/29

16.1

34.0

8.17

4

115a

705

oc

P

7/ 1/29

16.1

34.0

8.17

4

116

703

50

oc

n

7/ 1/29

10.6

33.8

8.11

23

116

704

100

oc

n

7/ 1/29

6.7

33.8

116

706

50

oc

P

7/ 1/29

10.6

33.8

8.11

23

116

707

100

r

P

7/ 1/29

6.7

33.8

. • .

117

708

r

n

7/ 3/29

15.9

34.3

8.17

3

117

709

50

oc

n

7/ 3/29

12.5

34.2

8.06

51

117

710

100

oc

n

7/ 3/29

8.8

34.1

7.98

84

117

711

c

P

7/ 3/29

15.9

34.3

8.17

3

117

712

50

oc

P

7/ 3/29

12.5

34.2

8.06

51

Table 15. Distributional and environmental records for Ceratium setaceum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

P04

mg/m3

70

70

110

434
435
666

50

100

r
r
oc

1/13/29
1/13/29
5/31/29

21.2
15.4
17.9

35.1
35.0
34.7

8.05
7.88
8.14

103

178

11

Table 16. Distributional and environmental records for Ceratium bigelowii

Station

Sample

Depth

Relative

Appa-

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/nH

45

257

50

r

n

11/19/28

22.4

35.2

8.13

46

47

267

50

r

n

11/23/28

23.8

36.0

8.23

20

47

268

100

r

n

11/23/28

22.7

36.2

8.23

20

48

272

50

r

n

11/25/28

23.6

36.4

8.24

16

49

278

50

r

n

11/27/28

22.6

36.1

8.26

13

54

321

100

r

n

12/14/28

18.7

35.4

8.16

20

56

330

100

r

n

12/18/28

16.6

34.8

8.11

12

57

337

100

r

n

12/20/28

14.3

34.4

8.10

40

78

477

100

r

n

2/20/29

21.9

36.2

8.14

34

78

478

r

p

2/20/29

24.6

36.0

8.17

32

81

492

50

r

n

2/26/29

26.4

35.9

8.19

38

89

529

50

r

n

3/23/29

28.6

35.8

8.27

12

89

530

r

p

3/23/29

28.4

35.6

8.25

21

89

532

r

n

3/23/29

(28.4

35.6

8.25

21)

95

563

50

oc

n

4/24/29

29.3

34.9

8.24

16

95

564

100

r

n

4/24/29

28.5

35.4

8.22

21

95

567

100

r

p

4/24/29

28.5

35.4

8.22

21

96

568

r

n

4/26/29

29.3

35.3

8.23

12

96

569

50

oc

n

4/26/29

29.2

35.3

8.23

12

96

570

100

r

n

4/26/29

28.2

35.7

8.19

25

100

596

50

r

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

r

n

5/ 4/29

27.6

34.7

8.22

12

103

615

r

n

5/11/29

26.0

35.0

8.25

S

105

627

50

r

n

5/15/29

26.8

34.9

8.23

5

106

634

50

r

n

5/17/29

27.0

35.0

8.23

5

107

639

r

n

5/19/29

28.0

34.4

8.23

5

107

641

100

r

n

5/19/29

26.8

34.9

8.23

11

108

648

100

r

n

5/27/29

25.2

35.0

8.23

4

109

655

50

r

n

5/29/29

23.1

35.0

8.22

3

136

829

100

r

n

9/16/29

18.6

35.0

8.39

3

68

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 16. Distributional and environmental records for Ceratium bigelowii- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

154

958

r

n

10/31/29

28.3

34.2

8.39

7

154

959

50

r

n

10/31/29

28.2

34.2

8.40

7

155

965

r

n

11/ 2/29

27.8

34.9

8.29

29

155

966

50

r

n

11/ 2/29

27.7

34.9

8.30

30

155

967

100

r

n

11/ 2/29

27.2

35.0

8.30

35

158

983

r

n

11/ 8/29

28.2

35.6

8.34

36

158

984

50

r

n

11/ 8/29

28.2

35.6

8.39

50

159

990

oc

n

11/11/29

28.6

35.7

8.37

15

159

991

50

oc

n

11/11/29

28.5

35.7

8.39

15

159

992

100

r

n

11/11/29

28.0

35.7

8.37

23

Table 17. Distributional and environmental records for Ceratium inflatum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

k/ *»*V*Xi^*«

(m)

abundance

ratus

(°C)

(0/00)

mg/m3

17

112

50

r

n

8/15/28

21.9

36.6

8.28

12

18

118

50

r

n

8/17/28

22.4

36.8

8.24

5

20-21

131

r

n

8/24/28

(26.6

36.3

8.32

4)

24

151

r

P

8/31/28

27.2

35.2

8.32

4

37

218

50

r

n

11/ 1/28

18.8

31.5

8.00

121

53

304

50

r

n

12/15/28

21.2

35.8

8.20

13

54

320

50

r

n

12/14/28

19.8

35.4

8.18

17

56

330

100

r

n

12/18/28

16.6

34.8

8.11

12

90

533

r

n

3/25/29

28.5

35.5

8.27

21

99

589

r

n

5/ 2/29

27.9

34.9

8.21

12

99

590

50

oc

n

5/ 2/29

27.8

34.9

8.22

12

99

595

r

n

5/ 2/29

27.9

34.9

8.21

12

100

597

100

r

n

5/ 4/29

27.6

34.7

8.22

12

101

603

r

n

5/ 7/29

26.3

34.7

8.24

8

101

605

100

r

n

5/ 7/29

25.2

35.1

8.23

8

104

621

r

n

5/13/29

25.8

35.2

8.24

7

104

625

50

r

p

5/13/29

25.8

35.2

8.24

7

IDS

627

50

r

n

5/15/29

26.8

34.9

8.23

5

IDS

629

100

r

n

5/15/29

25.2

35.1

8.23

5

106

633

r

n

5/17/29

27.2

35.0

8.23

5

106

634

50

r

n

5/17/29

27.0

35.0

8.23

5

106

635

100

r

n

5/17/29

25.6

35.1

8.23

5

107

642

r

p

5/19/29

28.0

34.4

8.23

5

111

672

50

r

P

6/ 3/29

19.4

34.6

8.17

5

138

845

100

r

n

9/20/29

22.2

34.8

8.31

3

142

874

100

r

n

10/ 8/29

16.6

34.4

8.27

7

145

890

r

n

10/13/29

22.3

34.6

8.29

6

145

894

50

r

P

10/13/29

18.7

34.3

8.34

6

151

942

50

r

P

10/26/29

18.3

34.4

158

983

r

n

11/ 8/29

28.2

35.6

8.34

36

159

990

r

n

11/11/29

28.6

35.7

8.37

15

Table 18. Distributional and environmental records for Ceratium longirostrum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

(m)

abimdance

ratus

(°C)

(0/00)

mg/m3

lb

8

100

r

n

5/18/28

(20.3

36.5

8.18

36)

20

128

r

n

8/22/28

26.0

36.6

8.37

5

49

277

r

n

11/27/28

23.4

36.2

8.27

13

49

279

100

r

n

11/27/28

21.6

35.9

8.26

13

49

282

100

r

P

11/27/28

21.6

35.9

8.26

13

49a

283

r

n

11/29/28

(23.2

36.0

8.23

13)

78

479

50

r

P

2/20/29

23.8

36.1

8.14

32

84

505

r

n

3/ 4/29

27.8

36.2

8.23

24

93

551

50

oc

n

3/31/29

28.5

34.8

8.30

28

93

553

100

r

P

3/31/29

27.6

35.8

8.27

29

96

570

100

r

n

4/26/29

28.2

35.7

8.19

25

96

573

100

r

P

4/26/29

28.2

35.7

8.19

25

98

585

50

r

p

4/30/29

26.9

35.3

8.16

28

101

605

100

oc

n

5/ 7/29

25.2

35.1

8.23

8

102

609

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

610

50

oc

n

5/ 9/29

25.8

35.0

8.24

8

103

615

oc

n

5/11/29

26.0

35.0

8.25

5

APPENDIX

69

Table 18. Distributional and environmental records for Ceratlum longirostrum

--Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4

mg/m^

103

616

50

r

n

5/11/29

25.8

35.2

8.25

5

103

617

100

r

n

5/11/29

24.8

35.2

8.25

5

103

618

r

P

5/11/29

26.0

35.0

8.25

5

104

621

r

n

5/13/29

26.1

35.2

8.24

7

106

635

100

r

n

5/17/29

25.6

35.1

8.23

5

107

639

r

n

5/19/29

28.0

34.4

8.23

5

107

642

r

P

5/19/29

28.0

34.4

8.23

5

145

890

r

n

10/13/29

22.3

34.6

8.29

6

145

893

r

P

10/13/29

22.3

34.6

8.29

6

Table 19. Distributional and environmental records for Ceratlum falcatum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/m3

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

50

286

100

r

n

11/29/28

20.5

35.7

8.22

13

50a

291

r

n

12/ 1/28

22.8

35.6

8.22

16

51

293

r

n

12/ 1/28

22.8

35.6

8.22

16

51

294

50

r

n

12/ 1/28

20.5

35.6

8.22

17

51

295

100

r

n

12/ 1/28

20.0

35.6

8.22

17

51

296

r

P

12/ 1/28

22.8

35.6

8.22

16

52

299

r

n

12/ 3/28

22.8

35.6

8.22

16

52

301

100

r

n

12/ 3/28

18.2

35.2

8.17

8

53

304

50

r

n

12/ 5/28

21.2

35.8

8.20

13

53-54

310

r

n

12/14/28

(23.0

35.6

8.22

11)

55

323

r

n

12/16/28

20.4

34.9

8.19

12

55

324

50

r

n

12/16/28

18.7

35.0

8.18

12

56

330

100

r

n

12/18/28

16.6

34.8

8.11

12

56

331

r

P

12/18/28

20.8

34.9

8.13

9

56-57

334

r

n

12/20/28

(19.0

34.5

8.14

20)

69

425

50

r

n

1/12/29

17.4

35.1

7.99

151

69

426

100

r

n

1/12/29

14.6

34.8

7.86

198

71

441

r

p

2/ 6/29

23.5

35.2

8.13

58

74

459

r

P

2/12/29

24.2

35.6

8.17

68

76

467

r

n

2/16/29

23.4

35.9

8.15

50

80

488

r

P

2/24/29

26.0

35.9

8.20

36

81

493

r

p

2/26/29

26.5

35.8

8.19

38

82

498

r

P

2/28/29

27.2

36.3

8.21

34

86

516

r

n

3/ 9/29

28.3

36.2

8.29

20

86

518

r

P

3/ 9/29

28.3

36.2

8.29

20

87

521

r

n

3/11/29

27.8

36.1

8.28

17

87

522

50

r

n

3/11/29

26.5

36.1

8.26

20

87

523

r

p

3/11/29

27.8

36.1

8.28

17

88

526

r

n

3/21/29

28.5

35.9

8.23

16

89

532

r

n

3/23/29

(28.4

35.6

8.25

21)

90

533

r

n

3/25/29

28.5

35.5

8.27

21

90

535

r

p

3/25/29

28.5

35.5

8.27

21

102

609

oc

n

5/ 9/29

25.8

35.0

8.24

8

108

697

50

r

n

5/27/29

26.8

35.0

8.24

4

109

656

100

r

n

5/29/29

19.4

34.8

8.18

5

110

664

r

p

5/31/29

23.9

34.7

8.18

5

138

843

r

n

9/20/29

26.1

34.8

8.35

5

138

844

50

r

p

9/20/29

25.6

34.7

8.30

3

139

850

50

r

n

9/22/29

25.8

34.9

8.31

6

140

858

100

r

n

10/ 3/29

25.5

35.0

8.34

7

149

920

r

n

10/21/29

23.5

35.0

8.34

6

149

927

r

p

10/21/29

23.5

35.0

8.34

6

150

930

50

r

n

10/23/29

22.8

34.8

8.35

10

150

931

100

r

n

10/23/29

19.6

34.6

8.32

11

152

945

50

r

n

10/27/29

14.2

34.5

7.87

53

158

985

100

r

n

10/27/29

27.6

35.9

8.39

48

70

CERATIUM m THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 20. Distributional and environmental records for Ceratium extensum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

1

1

r

n

5/12/28

24.0

36.2

8.16

34

la

5

r

n

5/18/28

(22.3

36.3

8.20

46

lb

6

oc

n

5/18/28

122.3

36.3

8.20

46

lb

7

50

oc

n

5/18/28

(22.3

36.3

8.20

46

2

13

100

oc

n

5/18/28

19.8

36.4

8.21

33

2

14

r

n

5/18/28

20.5

36.4

8.23

58

3

17

c

n

5/21/28

15.5

36.1

8.15

99

3

18

50

00

n

5/21/28

15.0

36.0

8.19

30

6b

44

c

n

5/31/28

(12.4

35.6

8.15

21)
21)

6b

45

.oc

n

5/31/28

(12.4

35.6

8.15

6b

46

oc

n

5/31/28

12.4

35.6

8.15

21)

6c

48

oc

n

5/31/28

12.4

35.6

8.15

21)

6c

49

c

n

5/31/28

(12.4

35.6

8.15

21)

16

103

r

P

8/13/28

25.9

36.2

8.24

8

16

105

100

r

P

8/13/28

19.9

36.5

8.17

13

16

106

r

n

8/13/28

25.9

36.2

8.24

8

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

17

109

r

P

8/15/28

26.2

36.6

8.29

9

18

114

r

P

8/17/28

27.0

37.0

8.23

5

19

123

50

r

n

8/20/28

25.2

37.1

8.27

5

31

193

100

r

n

10/ 3/28

23.4

36.5

8.19

28

33

197A

r

P

10/ 8/28

28.5

35.6

8.23

4

34

200A

oc

P

10/ 9/28

28.5

35.9

8.28

2

35-36

209

oc

n

10/26/28

(27.4

29.7

8.31

15)

45

256

r

n

11/19/28

22.4

35.3

8.12

38

45

257

50

r

n

11/19/28

22.4

35.2

8.13

46

45

258

100

r

n

11/19/28

18.6

35.1

8.00

50

45

259

r

P

11/19/28

22.4

35.3

8.12

38

46

261

oc

n

11/21/28

23.3

35.3

8.16

36

46

262

50

r

n

11/21/28

23.2

35.3

8.16

40

46

263

100

r

n

11/21/28

22.5

35.4

8.17

40

46

264

r

P

11/21/28

23.3

35.3

8.16

36

46

265

50

oc

P

11/21/28

23.2

35.3

8.16

40

47

266

r

n

11/23/28

23.9

36.0

8.23

17

47

267

50

oc

n

11/23/28

23.8

36.0

8.23

20

47

268

100

oc

n

lf/23/28

22.7

36.2

8.23

20

47

269

oc

P

11/23/28

23.9

36.0

8.23

17

47

270

50

r

p

11/23/28

23.8

36.0

8.23

20

48

272

50

r

n

11/25/28

23.6

36.4

8.24

16

48

273

100

oc

n

11/25/28

22.7

36.3

8.26

16

48

274

r

p

11/25/28

23.6

36.4

8.23

13

49

277

oc

n

11/27/28

23.4

36.2

8.27

13

49

279

100

oc

n

11/27/28

21.6

35.9

8.26

13

49

280

r

p

11/27/28

23.4

36.2

8.27

13

49

281

50

r

p

11/27/28

22.6

36.1

8.26

13

49a

283

oc

n

11/28/28

(23.2

36.0

8.23

13)

50

284

oc

n

11/29/28

23.2

36.0

8.23

13

50

286

100

oc

n

11/29/28

20.5

35.7

8.22

13

50-51

290

r

n

11/30/28

(22.8

35.6

8.22

16)

50-51

291

oc

n

11/30/28

122.8

35.6

8.22

16)

50-51

292

oc

n

11/30/28

22.8

35.6

8.22

16)

51

293

oc

n

12/ 1/28

(22.8

35.6

8.22

16)

51

294

50

r

n

12/ 1/28

20.5

35.6

8.22

17

51

295

100

r

n

12/ 1/28

20.0

35.6

8.22

17

51

296

oc

P

12/ 1/28

22.8

35.6

8.22

16

52

299

r

n

12/ 3/28

22.5

35.4

8.21

8

53-54

310

r

n

12/10/28

(23.0

35.6

8.22

11

53-54

311

r

n

12/10/28

23.0

35.6

8.22

11

53-54

312

oc

n

12/10/28

(23.0

35.6

8.22

11

54

319

r

n

12/14/28

23.4

35.5

8.22

9

54

320

50

oc

n

12/14/28

19.8

35.4

8.18

17

54

322

r

P

12/14/28

23.4

35.5

8.22

9

55

323

r

n

12/16/28

20.4

34.9

8.19

12

56

330

100

r

n

12/18/28

16.6

34.8

8.11

12

56

333

100

r

P

12/18/28

16.6

34.8

8.11

12

57

335

oc

n

12/20/28

19.0

34.5

8.14

20

59

351

r

P

12/24/28

16.3

34.0

8.10

38

60

355

50

r

n

12/26/28

13.4

34.0

8.06

54

60

356

100

r

n

12/26/28

10.6

34.0

8.03

62

61

368

100

r

P

12/28/28

10.8

34.0

8.03

80

61-62

369

r

n

12/28/28

(16.9
16.9

34.0

8.05

46)
46

61-62

370

r

n

12/28/28

34.0

8.05

63

381

100

r

n

1/ 1/29

15.6

34.6

8.08

24

63-64

386

r

n

1/ 1/29

(20.5

34.6

8.07

21

63-64

387

r

n

1/ 2/29

20.5

34.6

8.07

21

63-64

388

r

n

1/ 2/29

20.5

34.6

8.07

21

APPENDIX

71

Table 20. Distributional and environmental records for Ceratlum extensum- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

mg/m3

(m)

abundance

ratus

(°C)

(o/oo)

63-64

389

r

n

1/ 2/29

20.5

34.6

8.07

21)

63-64

390

r

n

1/ 2/29

20.5

34.6

8.07

21)

63-64

391

r

n

1/ 3/29

20.5

34.6

8.07

21)

63-64

392

r

n

1/ 3/29

20.5

34.6

8.07

21)

63-64

393

r

n

1/ 3/29

20.5

34.6

8.07

21)

64

394

r

n

1/ 3/29

20.6

34.6

8.12

21

64

398

r

p

1/ 3/29

20.6

34.6

8.12

21

64-65

402

r

n

1/ 4/29

(20.6

34.6

8.12

21)

65

408

r

P

1/ 5/29

20.2

34.5

8.10

24

67

416

DC

n

1/ 8/29

19.3

34.9

8.11

21

68

422

r

P

1/10/29

19.2

35.1

8.14

29

76

468

50

r

n

2/16/29

22.1

35.9

8.14

42

76

469

100

r

n

2/16/29

21.2

35.8

8.12

45

77

472

r

n

2/18/29

23.7

36.0

8.19

16

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

478

r

P

2/20/29

24.6

36.0

8.17

32

78

479

SO

r

P

2/20/29

23.8

36.1

8.14

32

78

480

100

r

p

2/20/29

21.9

36.2

8.14

34

79

481

oc

n

2/22/29

25.2

36.0

8.17

34

79

482

50

oc

n

2/22/29

24.5

36.1

8.17

34

79

483

100

r

n

2/22/29

21.8

36.2

8.13

45

79

484

r

p

2/22/29

25.2

36.0

8.17

34

80

486

r

n

2/24/29

26.0

35.9

8.20

36

80

488

r

P

2/24/29

26.0

35.9

8.20

36

80

489

50

oc

P

2/24/29

25.9

36.0

8.19

29

80

490

100

r

P

2/24/29

23.4

36.2

8.16

32

81

491

r

n

2/26/29

26.5

35.8

8.19

38

81

492

50

c

n

2/26/29

26.4

35.9

8.19

38

81

493

oc

P

2/26/29

26.5

35.8

8.19

38

81

494

50

oc

P

2/26/20

26.4

35.9

8.19

38

81

495

100

r

p

2/26/29

23.6

36.2

8.18

36

82

496

.

oc

n

2/28/29

27.2

36.3

8.21

34

82

497

50

oc

n

2/28/29

27.2

36.3

8.21

34

82

498

oc

P

2/28/29

27.2

36.3

8.21

34

83

501

c

n

3/ 2/29

27.5

36.3

8.24

29

83

502

50

r

n

3/ 2/2S

27.4

36.5

8.24

25

83

504

50

r

P

3/ 2/29

27.4

36.5

8.24

25

84

505

oc

n

3/ 4/29

27.8

36.2

8.23

24

84

506

50

oc

n

3/ 4/29

27.5

36.4

8.21

24

85

510

oc

n

3/ 6/29

27.9

36.2

8.22

40

85

511

50

r

n

3/ 6/29

27.8

36.2

8.22

40

85

512

r

p

3/ 6/29

27.9

36.2

8.22

40

86

516

oc

n

3/ 9/29

28.3

36.2

8.29

20

86

517

50

oc

n

3/ 9/29

27.4

36.2

8.29

17

86

518

r

p

3/ 9/29

28.3

36.2

8.29

20

87

521

oc

n

3/11/29

27.8

36.1

8.28

17

87

523

r

P

3/11/29

27.8

36.1

8.28

17

87

525

100

r

P

3/11/29

23.9

36.0

8.23

20

88

526

oc

n

3/21/29

28.5

35.9

8.23

16

89

528

oc

n

3/23/29

28.4

35.6

8.25

21

89

529

50

oc

n

3/23/29

28.6

35.8

8.27

12

89

530

r

P

3/23/29

28.4

35.6

8.25

21

89

532

r

n

3/23/29

(28.4

35.6

8.25

21)

90

533

c

n

3/25/29

28.5

35.5

8.27

21

90

535

oc

P

3/25/29

28.5

35.5

8.27

21

90

536

50

oc

P

3/25/29

28.6

35.6

8.26

21

91

540

c

n

3/27/29

28.7

35.1

8.30

21

91

541

50

c

n

3/27/29

28.5

35.2

8.30

24

91

542

oc

p

3/27/29

28.7

35.1

8.30

21

91

543

50

r

p

3/27/29

28.5

35.2

8.30

24

91

544

100

r

p

3/27/29

25.8

36.0

8.25

30

92

545

oc

n

3/29/29

28.5

35.3

8.29

28

92

546

50

c

n

3/29/29

28.4

35.4

8.29

28

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

551

50

c

n

3/31/29

28.5

34.8

8.30

28

93

552

r

P

3/31/29

28.7

34.7

8.30

28

93

553

100

r

p

3/31/29

27.6

35.8

8.27

29

94

558

50

r

n

4/22/29

29.3

34.7

8.25

14

94

559

100

r

n

4/22/29

28.5

35.6

8.21

25

94

561

50

r

P

4/22/29

29.3

34.7

8.25

14

95

565

■ r

P

4/24/29

29.4

34.7

8.26

14

95

568A

oc

n

4/25/29

(29.4

34.7

8.26

14)

96

569

50

oc

n

4/26/29

29.2

35.3

8.23

12

96

572

50

r

P

4/26/29

29.2

35.3

8.23

12

97

577

r

P

4/28/29

28.3

35.2

8.16

24

72

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 20. Distributional and environmental records for Ceratium extensum — Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

97

578

50

r

P

4/28/29

28.0

35.4

8.16

21

98

584

r

p

4/30/29

27.0

35.3

8.16

24

98

585

50

r

p

4/30/29

26.9

35.3

8.16

28

99

594

100

r

P

5/ 2/29

27.8

35.0

8.22

17

99

595

r

n

5/ 2/29

27.9

34.9

8.21

12

100

596

50

r

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

r

n

5/ 4/29

27.6

34.7

8.22

12

100

598

r

p

5/ 4/29

27.7

34.7

8.21

10

102

609

c

n

5/ 9/29

25.8

35.0

8.24

8

102

610

50

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

611

100

oc

n

5/ 9/29

25.6

35.0

8.23

8

102

612

r

p

5/ 9/29

25.8

35.0

8.24

8

102

613

50

r

p

5/ 9/29

25.8

35.0

8.24

8

103

615

oc

n

5/11/29

26.0

35.0

8.25

5

103

616

50

r

n

5/11/29

25.8

35.2

8.25

5

103

617

100

oc

n

5/11/29

24.8

35.2

8.25

5

103

618

r

P

5/11/29

26.0

35.0

8.25

5

104

622

50

oc

n

5/13/29

25.8

35.2

8.24

7

104

626

100

r

P

5/13/29

25.3

35.3

8.21

7

105

627

50

c

n

5/15/29

26.8

34.9

8.23

5

105

628

50

c

n

5/15/29

26.8

34.9

8.23

5

105

630

oc

p

5/15/29

26.9

34.9

8.23

5

105

631

50

oc

P

5/15/29

26.8

34.9

8.23

5

106

633

c

n

5/17/29

27.2

35.0

8.23

5

106

634

50

oc

n

5/17/29

27.0

35.0

8.23

5

106

635

100

c

n

5/17/29

25.6

35.1

8.23

5

106

636

r

P

5/17/29

27.2

35.0

8.23

5

106

637

50

r

P

5/17/29

27.0

35.0

8.23

5

107

639

c

n

5/19/29

28.0

34.4

8.23

5

107

640

50

oc

n

5/19/29

27.9

34.4

8.23

4

107

641

100

oc

n

5/19/29

26.8

34.9

8.23

11

107

642

r

P

5/19/29

28.0

34.4

8.23

5

107

643

50

r

P

5/19/29

27.9

34.4

8.23

4

107

645

50

oc

P

5/19/29

27.9

34.4

8.23

•4

108

649

r

p

5/27/29

28.4

35.0

8.25

4

Guam

652

oc

n

5/ /29

...

109

654

oc

n

5/29/29

27; 4

35.0

8.23

4

109

655

50

r

n

5/29/29

23.1

35.0

8.22

3

109

658

50

oc

p

5/29/29

23.1

35.0

8.22

3

110

664

oc

P

5/31/29

23.9

34.7

8.18

5

112

675

50

r

n

6/ 5/29

21.7

34.6

8.23

7

112

677

r

P

6/ 5/29

23.2

34.6

8.22

7

128

779

r

n

7/25/29

16.4

33.0

8.12

29

129

780

r

n

7/27/29

16.3

33.1

8.13

25

130

787

oc

n

9/ 4/29

16.2

33.4

8.34

36

131

793

r

P

9/ 6/29

19.3

33.4

8.34

.••>

132

801

r

P

9/ 8/29

21.0

33.9

8.34

15

133

814

100

r

P

9/10/29

18.4

34.8

8.31

7

134

808

r

n

9/12/29

22.9

34.7

8.34

6

134

809

50

r

n

9/12/29

19.8

34.6

8.34

6

137

840

r

P

9/18/29

25.5

35.0

8.39

4

140

862

100

r

p

10/ 3/29

25.5

35.0

8.34

7

141

864

r

n

10/ 5/29

25.9

35.2

8.34

5

141

868

r

p

10/ 5/29

25.9

35.2

8.34

5

142

872

r

n

10/ 7/29

24.1

34.8

8.33

5

142

873

50

oc

n

10/ 7/29

21.8

34.8

8.30

5

142

876

r

p

10/ 7/29

24.1

34.8

8.33

5

142

877

50

r

P

10/ 7/29

21.8

34.8

8.30

5

142

879

100

r

P

10/ 7/29

16.6

34.4

8.27

7

143

881

oc

n

10/ 9/29

22.4

34.4

8.30

6

143

882

50

oc

n

10/ 9/29

19.0

34.2

8.34

6

143

883

100

oc

n

10/ 9/29

13.8

34.1

8.30

10

143

884

r

p

10/ 9/29

19.0

34.2

8.34

6

143

885

50

oc

n

10/ 9/29

19.0

34.2

8.34

6

144

886

r

n

10/11/29

23.3

35.0

8.37

6

144

887

oc

P

10/11/29

23.3

35.0

8.37

6

144

888

50

r

P

10/11/29

21.1

34.7

8.33

6

144

889

100

r

P

10/11/29

16.6

34.5

8.37

6

145

890

oc

n

10/13/29

22.3

34.6

8.29

6

145

891

50

oc

n

10/13/29

18.7

34.3

8.34

6

145

893

oc

P

10/13/29

22.3

34.6

8.29

6

145

894

50

oc

P

10/13/29

18.7

34.3

8.34

6

146

898

100

r

n

10/15/29

19.7

34.3

8.26

7

146

900

r

P

10/15/29

22.4

34.9

8.37

6

146

902

100

r

P

10/15/29

19.7

34.3

8.26

7

APPENDIX

73

Table 20. Distributional and environmental records for Ceratlum extensum - -Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature

Cc)

Salinity
(o/oo)

PH

PO4
mg/m3

152

946

100

r

n

10/27/29

11.4

34.7

7.76

75

152

949

50

r

P

10/27/29

14.2

34.5

7.87

53

158

983

oc

n

11/ 8/29

28.2

35.6

8.34

36

158

984

50

oc

n

11/ 8/29

28.2

35.6

8.39

50

158

985

100

c

n

11/ 8/29

27.6

35.9

8.39

48

159

991

50

oc

n

11/11/29

28.5

35.7

8.39

15

159

992

100

r

n

11/11/29

28.0

35.7

8.37

23

Table 21. Distributional and environmental records for Ceratium fusus

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

CC)

(0/00)

mg/m3

3

19

100

r

n

5/21/28

13.6

35.9

8.10

48

3

20

c

n

5/21/28

15.5

36.1

8.15

99

3-4

21

r

n

5/23/28

(15.1

36.0

8.15

96)

4

24

50

r

n

5/23/28

14.4

36.0

8.15

21

4

25

r

n

5/23/28

14.6

35.9

8.16

92

5

28

50

oc

n

5/25/28

14.6

36.0

8.21

36

5

29

oc

n

5/25/28

15.0

35.9

8.19

16

5a

33

c

n

5/31/28

12.4
12.4

35.6

8.15

21)
21)

5a

34

r

n

5/31/28

35.6

8.15

6

35

c

n

5/31/28

12.4

35.6

8.15

21

6

36

50

c

n

5/31/28

11.6

35.6

8.12

32

6

37

100

c

n

5/31/28

11.3

35.6

8.08

41

6

42

c

n

5/31/28

(12.4

35.6

8.15

21)

6b

43

oc

n

5/31/28

112.4

35.6

8.15

21)

6b

44

c

n

5/31/28

12.4

35.6

8.15

21

6b

45

oc

n

5/31/28

12.4

35.6

8.15

21

6b

46

oc

n

5/31/28

12.4

35.6

8.15

21

6c

48

oc

n

5/31/28

12.4

35.6

8.15

21)

6c

49

c

n

5/31/28

12.4

35.6

8.15

21)

7

56

oc

n

7/13/28

8.9

35.2

8.08

34

8

59

oc

n

7/15/28

10.3

35.2

7.93

13

8

60

50

r

n

7/15/28

9.1

35.2

7.95

27

8

61

100

oc

n

7/15/28

8.4

35.3

7.95

54

9

64

a

n

7/28/28

11.2

35.1

8.08

20

9

65

50

a

n

7/28/28

8.4

35.1

7.96

55

9

66

100

a

n

7/28/28

7.6

35.1

7.98

56

10

68

a

n

7/30/28

10.9

34.9

8.08

28

10

69

50

a

n

7/30/28

10.0

34.9

8.04

34

11

73

oc

n

8/ 1/28

10.7

34,9

8.06

27

11

74

50

oc

n

8/ 1/28

7.3

34.9

7.92

63

11

75

r

p

8/ 1/28

10.7

34.9

8.06

27

11

76

50

r

P

8/ 1/28

7.3

34.9

7.92

63

11

77

100

r

P

8/ 1/28

6.3

35.1

7.90

66

11a

78

a

n

8/ 1/28

(10.7

34.9

8.06

27)

13

83

r

n

8/ 7/28

11.3

32.7

8.09

19

13a

89

r

n

8/ 9/28

(21.2

35.2

8,18

11)

14

93

c

P

8/ 9/28

21.2

35.2

8.18

11

14

95

50

r

n

8/ 9/28

15.0

35.1

8.18

16

15

97

oc

P

8/11/28

34.8

36.4

8.21

11

15

98

50

oc

p

8/11/28

19.8

36.5

8.21

8

15

99

100

oc

p

8/11/28

18.4

36.4

8.20

19

15

100

oc

n

8/11/28

24.8

36.4

8.21

11

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

16

103

r

p

8/13/28

25.9

36.2

8.24

8

16

104

50

r

P

8/13/28

24.4

36.4

8.23

8

16

105

100

r

P

8/13/28

1S.9

36.5

8.17

13

16

106

r

n

8/13/28

25.9

36.2

8.24

8

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

16

108

100

oc

n

8/13/28

19.9

36.5

8.17

13

17

109

r

P

8/15/28

26.2

36.6

8.29

9

17

110

50

oc

P

8/15/28

21.9

36.6

8.28

12

17

111

a

n

8/15/28

26.2

36.6

8.29

9

18

114

oc

P

8/17/28

27.0

37.0

8.23

5

18

117

r

n

8/17/28

27.0

37.0

8.23

5

22

139

r

P

8/27/28

26.7

36.0

8.26

8

23

145

r

P

8/29/28

27.2

35.9

8.25

4

25

158

50

r

P

9/ 3/28

21.5

36.0

8.22

12

33

197A

r

p

10/ 8/28

28.5

35.6

8.23

4

35

207

50

r

P

10/26/28

16.8

34.7

7.92

138

37

219

100

r

n

11/ 1/28

15.1

34.9

7.82

153

74

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 21. Distributional and environmental records for Ceratium fusus- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

CC)

(o/oo)

mg/m3

40

232

r

P

11/ 8/28

22.2

33.7

8.21

24

41

236

50

r

n

11/10/28

14.6

35.0

7.94

58

41a

240

r

n

11/13/28

(18.7

34.7

8.06

45)

45

256

r

n

11/19/28

22.4

35.3

8.12

38

45

258

100

r

n

11/19/28

18.6

35.1

8.00

50

45

259

CO

P

11/19/28

22.4

35.3

8.12

38

45

260

50-0

DC

P

11/19/28

22.4

35.2

8.13

46

47

267

50

r

n

11/23/28

23.8

36.0

8.23

20

50

285

50

r

n

11/29/28

22.0

35.9

8.23

13

50

286

100

r

n

11/29/28

20.5

35.7

8.22

13

50

289

100

r

P

11/29/28

20.5

35.7

8.22

13

50-51

292

r

n

12/ 1/28

(22.8

35.6

8.22

16)

51

293

r

n

12/ 1/28

22.8

35.6

8.22

16

51

295

100

r

n

12/ 1/28

20.0

35.6

8.22

17

58

341

DC

n

12/22/28

17.0

34.0

8.12

20

58

342

50

OC

n

12/22/28

14.8

34.0

8.12

25

58

344

r

P

12/22/28

17.0

34.0

8.12

20

58

345

SO

OC

P

12/22/28

14.8

34.0

8.12

25

59

348

r

n

12/24/28

16.3

34.0

8.10

38

59

350

100

oc

n

12/24/28

11.4

34.1

8.03

72

59

351

OC

P

12/24/28

16.3

34.0

8.10

38

59

352

50

r

P

12/24/28

14.0

34.0

8.08

38

60

355

50

oc

n

12/26/28

13.4

34.0

8.06

54

60

356

100

OC

n

12/26/28

10.6

34.0

8.03

62

60

358

50

r

P

12/26/28

13.4

34.0

8.06

54

60

359

r

P

12/26/28

15.0

34.0

8.07

50

60-61

362

r

n

12/26/28

(15.0

34.0

8.07

50)

61

368

100

r

P

12/28/28

10.8

34.0

8.03

80

61-62

370

r

n

12/28/28

(16.9

34.0

8.05

46)

62

373

100

r

n

12/30/28

13.1

34.2

8.06

48

63-64

392

r

n

1/ 1/29

(20.5

34.6

8.07

21)

64

395

50

r

n

1/ 3/29

17.2

34.6

8.12

29

64-65

401

r

n

1/ 3/29

(20.6

34.6

8.12

21)

64-65

402

OC

n

1/ 3/29

20.6

34.6

8.12

21

64-65

403

r

n

1/ 3/29

(20.6

34.6

8.12

21)

65

408

r

P

1/ 5/29

20.2

34.5

8.10

24

65

409

50

r

P

1/ 5/29

16.5

34.5

8.10

25

67

416

OC

n

1/ 8/29

19.3

34.9

8.11

21

68

422

r

P

1/10/29

19.2

35.1

8.14

29

68

423

50

r

P

1/10/29

18.2

35.0

8.14

29

69

427

r

P

1/12/29

21.1

35.2

8.12

62

70

434

r

P

1/13/29

21.2

35.1

8.05

103

71

439

50

r

n

2/ 6/29

16.7

35.1

7.90

150

71

440

100

r

n

2/ 6/29

13.9

35.0

7.71

220

71

441

OC

P

2/ 6/29

23.5

35.2

8.13

58

71

442

50

r

P

2/ 6/29

16.7

35.1

7.90

150

72

444

50

r

n

2/ 8/29

18.7

35.4

8.12

60

72

446

r

P

2/ 8/29

24.9

35.3

8.16

50

72

448

100

r

P

2/ 8/29

14.8

35.0

7.94

154

74

459

OC

P

2/12/29

24.2

35.6

8.17

68

75

466

50

r

P

2/14/29

20.0

35.5

8.14

46

76

467

r

n

2/16/29

23.4

35.9

8.15

50

76

471

50

r

P

2/16/29

22.1

35.9

8.14

42

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

r

n

2/20/29

21.9

36.2

8.1')

34

78

478

r

P

2/20/29

24.6

36.0

8.17

32

78

479

50

OC

P

2/20/29

23.8

36.1

8.14

32

78

480

100

r

P

2/20/29

21.9

36.2

8.1''

34

79

483

100

r

n

2/22/29

21.8

36.2

8.13

45

80

488

OC

P

2/24/29

26.0

35.9

8.30

36

80

489

50

OC

P

2/24/29

25.9

36.0

8.19

29

80

490

100

OC

P

2/24/29

23.4

36.2

8.16

32

81

493

OC

P

2/26/29

26.5

35.8

8.19

38

81

494

50

r

P

2/26/29

26.4

35.9

8.19

38

81

495

100

r

P

2/26/29

23.6

36.2

8.18

36

82

"98

OC

P

2/28/29

27.2

36.3

8.21

34

82

500

100

r

P

2/28/29

24.4

36.5

8.19

34

85

513

50

OC

P

3/ 6/29

(27.8

36.2

8.22

40

85

514

100

r

P

3/ 6/29

25.2

36.3

8.23

42

86

519

50

r

P

3/ 9/29

27.4

35.2

8.29

17

89

530

r

p

3/23/29

28.4

35.6

8.25

21

90

536

50

r

P

3/25/29

28.6

35.6

8.26

21

92

549

100

r

P

3/29/29

26.2

36.0

8.28

28

93

551

50

OC

n

3/31/29

28.5

34.8

8.30

28

93

552

r

P

3/31/29

28.7

34.7

8.30

28

93

553

100

OC

P

3/31/29

27.6

35.8

8.27

29

APPENDIX

75

Table 21. Distributional and environmental records for Ceratium fusus- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/m3

Samoa

555

r

n

4/ /29

94

561

50

r

P

4/22/29

29! 3

34!?

8.'25

14

95

566

50

r

P

4/24/29

29.3

34.9

8.24

16

95

566

50

DC

P

4/24/29

29.3

34.9

8.24

16

95

568A

OC

n

4/24/29

(29.4

34.7

8.26

14)

96

572

50

DC

P

4/26/29

29.2

35.3

8.23

12

98

584

OC

P

4/30/29

27.0

35.3

8.16

24

100

602

100

r

P

5/ 4/29

27.6

34.7

8.22

12

110

666

100

r

P

5/31/29

17.9

34.7

8.14

11

111

668

r

n

6/ 3/29

20.1

34.5

8.18

5

111

669

50

DC

n

6/ 3/29

19.4

34.6

8.17

5

111

671

OC

P

6/ 3/29

20.1

34.5

8.18

5

111

672

50

OC

P

6/ 3/29

19.4

34.6

8.17

5

112

677

r

p

6/ 5/29

23.2

34.6

8.22

7

114

688

100

r

P

6/27/29

13.0

34.5

8.00

91

114

689

OC

P

6/27/29

19.9

34.3

8.15

7

115

692

OC

P

6/29/29

20.6

34.6

8.19

4

115

693

50

r

n

6/29/29

17.5

34.6

8.12

17

115

698

100

r

p

6/29/29

15.6

34.6

8.08

27

115a

700

r

n

7/ 1/29

(18.3

34.3

8.18

4)

116

706

50

r

p

7/ 1/29

10.6

33.8

8.11

23

120

726

OC

p

7/ 9/29

7.2

33.0

7.98

137

120

727

50

r

n

7/ 9/29

2.2

33.1

7.90

177

120

729

r

p

7/ 9/29

7.2

33.0

7.98

137

120

730

50

OC

p

7/ 9/29

2.2

33.1

7.90

177

121

732

c

n

7/11/29

7.5

32.9

7.98

141

121

733

50

r

n

7/11/29

3.6

33.1

7.92

159

121

735

OC

P

7/11/29

7.5

32.9

7.98

141

121

734

100

r

n

7/11/29

2.1

33.2

7.86

184

121

736

50

r

p

7/11/29

3.6

33.1

7.92

159

121

737

100

r

P

7/11/29

2.1

33.2

7.86

184

122

738

C

n

7/13/29

8.2

32.8

7.98

130

122

740

100

r

n

7/13/29

2.4

33.1

7.90

161

122

741

OC

P

7/13/29

8.2

32.8

7.98

130

123

744

OC

n

7/15/29

8.1

32.8

8.03

113

123

745

50

OC

n

7/15/29

4.4

32.9

7.94

150

123

746

100

OC

n

7/15/29

3.0

33.4

7.79

209

123

747

OC

P

7/15/29

8.1

32.8

8.03

113

124

749

OC

n

7/17/29

9.3

32.7

8.04

103

124

752

r

P

7/17/29

9.3

32.7

8.04

103

125

755

OC

n

7/19/29

10.5

32.8

8.03

125

125

756

50

OC

n

7/19/29

5.5

32.8

7.98

138

125

757

100

C

n

7/19/29

4.2

32.9

7.90

175

125

758

OC

p

7/19/29

10.5

32.8

8.03

125

125

759

50

OC

p

7/19/29

5.5

32.8

7.98

138

127

772

100

r

P

7/23/29

8.2

32.8

8.00

72

128

774

50

OC

p^

7/25/29

11.8

33.1

8.11

29

128

775

100

r

P*

7/25/29

10.2

33.2

8.06

46

128

777

50

OC

P

7/25/29

11.8

33.1

8.11

29

130

787

OC

n

9/ 4/29

16.2

33.4

8.34

36

131

795

r

P

9/ 6/29

19.3

33.4

8.34

141

869

50

r

p

10/ 5/29

24.8

35.3

8.34

"5

141

870

50

r

p

10/ 5/29

24.8

35.3

8.34

5

142

874

100

r

n

10/ 7/29

16.6

34.4

8.27

7

143

883

100

OC

n

10/ 9/29

13.8

34.1

8.30

10

144

887

r

p

10/11/29

23.3

35.0

8.37

6

144

889

100

r

p

10/11/29

16.6

34.5

8.37

6

151

941

r

P

10/26/29

26.0

34.0

Table 22. Distributional and environmental records for Ceratium tripos subsp. atlanticum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4
mg/m3

(m)

abundance

ratus

(°C)

(0/00)

1

1

oc

n

5/12/28

24.0

36.2

8.16

34

1

2

70

r

n

5/12/28

22.4

36.4

8.15

36)

la

3

oc

n

5/18/28

22.3

36.3

8.20

46

lb

7

50

c

n

5/18/28

21.7

36.3

8.20

38)

2

12

50

oc

n

5/18/28

20.5

36.4

8.21

46

2

13

100

c

n

5/18/28

19.8

36.4

8.21

46

2

14

c

n

5/18/28

20.5

36.4

8.23

58

2

15

r

n

5/18/28

20.5

36.4

8.23

58

3

17

oc

n

5/21/28

15.5

36.1

8.15

99

76

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 22. Distributional and environmental records for Ceratium tripos subsp. atlanticvun- -Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
CC)

Salinity
(o/oo)

pH

PO4
mg/m3

3

19

100

4

23

100

4

24

50

5a

33

5a

34

6

35

6

36

50

6

37

100

6a

40

6a

42

6b

43

6b

4"

6b

45

6b

46

6c

47

100

6c

48

6c

49

6e

52

6f

53

6g

5"

6h

55

7

56

7

57

50

7

58

100

8

59

8

60

50

8

61

100

9

64

9

65

50

9

66

100

9

67

10

68

10

69

50

10

70

100

11

73

11

74

50

11

75

11

77

100

11a

78

14

90

14

91

100

14

93

16

106

16

107

50

25

157

30

186

50

30

189

50

32

194

32

196 A

100

33

198A

50

37

217

37

218

50

37

220

39

225

39

226

50

39

227

100

41

235

42

241

42

242

50

43

246

100

43

247

44

253

45

256

45

258

100

60

354

60

355

50

60

356

100

60

357

60-61

360

60-61

361

60-61

362

61

363

61

364

50

61

365

100

61

366

61

367

50

61

368

100

r

r

r

c

r

c

c

c

r

c

c

c

c

c

c

c

c

r

c

c

oc

oc

oc

oc

c

c

oc

a

a

c

r

a

c

oc

oc

r

oc

r

oc

oc

oc

c

oc

c

r

r

r

oc

r

oc

oc

oc

oc

r

r

r

r

r

r

r

r

r

oc

oc

c

oc

oc

oc

c

oc

c

a

c

oc

a

c

c

n

5/21/28

13.6

n

5/23/28

13.4

n

5/23/28

14.4

n

5/31/28

(12.4

n

5/31/28

12.4

n

5/31/28

12.4

n

5/31/28

11.6

n

5/31/28

11.3

n

5/31/28

(12.4

n

6/ 2/28

(12.4

n

6/ 2/28

12.4

n

6/ 2/28

(12.4

n

6/ 2/28

12.4

n

6/ 2/28

(12.4

n

6/ 2/28

(11.3

n

6/ 2/28

1I2.4

n

6/ 4/28

12.4

n

7/ 8/28

(10.6

n

7/ 9/28

10.6

n

7/10/28

10.6

n

7/11/28

10.6

n

7/11/28

8.9

n

7/11/28

8.2

n

7/11/28

8.1

n

7/15/28

10.3

n

7/15/28

9.1

n

7/15/28

8.4

n

7/28/28

11.2

n

7/28/28

8.4

n

7/28/28

7.6

n

7/28/28

11.2

n

7/30/28

10.9

n

7/30/28

10.0

n

7/30/28

6.6

n

8/ 1/28

10.7

n

8/ 1/28

7.3

P

8/ 1/28

10.7

P

8/ 1/28

6.3

n

8/ 1/28

(10.7

n

8/ 9/28

21.2

P

8/ 9/28

14.0

P

8/ 9/28

21.2

n

8/13/28

25.9

n

8/13/28

24.4

P

9/ 3/28

27.5

P

9/15/28

27.8

n

9/15/28

27.8

n

10/ V28

28.0

P

10/ 5/28

22.2

P

10/ 8/28

28.2

n

11/ 1/28

27.1

n

11/ 1/28

18.8

P

11/ 1/28

27.1

n

11/ 1/28

24.8

n

11/ 6/28

16.3

n

11/ 6/28

14.0

n

11/10/28

20.4

n

11/13/28

18.7

n

11/13/28

17.2

n

11/15/28

13.6

P

11/15/28

19.6

P

11/17/28

20.7

n

11/19/28

22.4

n

11/19/28

18.6

n

12/26/28

15.0

n

12/26/28

13.4

n

12/26/28

10.6

P

12/26/28

15.0

n

12/26/28

15.0

n

12/26/28

115.0

n

12/26/28

15.0

n

12/28/28

16.9

n

12/28/28

14.0

n

12/28/28

10.8

P

12/28/28

16.9

P

12/28/28

14.0

P

12/28/28

10.8

35.9

8.10

48

35.9

8.12

50

36.0

8.15

21

35.6

8.15

21)

35.6

8.15

21

35.6

8.15

21

35.6

8.12

32

35.6

8.08

41

35.6

8.15

21)

35.6

8.15

21

35.6

8.15

21

35.6

8.15

21

35.6

8.15

21

35.6

8.15

21

35.6

8.08

41)

35.6

8.15

21)

35.6

8.15

21)

35.4

8.11

28

35.4

8.11

28)

35.4

8.11

28
28

35.4

8.11

35.2

8.08

34

35.2

8.03

47

35.2

8.04

57

35.2

7.93

13

35.2

7.95

27

35.3

7.95

54

35.1

8.08

20

35.1

7.96

55

35.1

7.98

56

35.1

8.08

20

34.9

8.08

28

34.9

8.04

34

35.0

7.95

52

34.9

8.06

27

34.9

7.92

63

34.9

8.06

27

35.1

7.90

66

34.9

8.06

27)

35.2

8.18

27

35.6

8.06

34

35.2

8.18

34

36.2

8.24

8

36.4

8.23

8

35.6

8.31

5

36.1

8.29

3

36.1

8.29

3

36.0

8.23

2

36.4

8.10

30

36.2

8.24

4

31.7

8.28

15

34.5

8.00

121

31.7

8.28

15

33.0

8.24

16

34.6

7.92

48

35.0

7.88

181

34.2

8.11

32

34.7

8.06

45

34.9

7.99

68

35.0

7.90

92

34.8

8.09

52

34.9

8.03

38

35.3

8.12

38

35.1

8.00

50

34.0

8.07

50

34.0

8.06

34

34.0

8.03

62

34.0

8.07

50

34.0

8.07

50)

34.0

8.07

50 i

34.0

8.07

50)

34.0

8.05

46

34.0

8.05

60

34.0

8.03

80

34.0

8.05

46

34.0

8.05

60

34.0

8.03

80

APPENDIX

77

Table 22. Distributional and environmental records for Ceratium tripos subsp. atlanticur- —Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
CO

Salinity
(o/oo)

pH

PO4

mg/m^

61-62

369

c

n

12/28/28

(16.9

34.0

8.05

46)

62

371

c

n

12/30/28

19.2

34.2

8.12

32

.62

372

50

c

n

12/30/28

16.2

34.3

8.10

28

62

373

100

c

n

12/30/28

13.1

34.2

8.06

48

62

375

50

oc

P

12/30/28

16.2

34.3

8.10

28

62-63

378

r

n

1/ 1/29

(16.2
(20.5

34.3

8.10

28)

63-64

391

r

n

1/ 1/29

34.6

8.07

2li

73

450

c

n

2/10/29

25.3

35.4

8.21

44

73

451

50

oc

n

2/10/29

18.7

35.4

8.05

122

73

452

100

oc

n

2/10/29

14.7

35.0

7.80

178

73

453

oc

P

2/10/29

25.3

35.4

8.21

44

74

457

50

r

n

2/12/29

19.2

35.4

8.06

80

79

481

r

n

2/22/29

25.2

36.0

8.17

34

79

482

50

oc

n

2/22/29

24.5

36.1

8.17

34

114

688

100

r

P

6/27/29

13.0

34.5

8.00

91

114

690

50

oc

P

6/27/29

16.2

34.6

8.04

63

114

691

100

r

n

6/27/29

13.0

34.5

8.00

91

115

692

oc

n

6/29/29

20.6

34.6

8.19

4

115

695-6

oc

P

6/29/29

20.6

34.6

8.19

4

115a

699

a

n

7/ 1/29

(18.3
(18.3

34.3

8.18

4)

115a

700

a

n

7/ 1/29

34.3

8.18

4)

115a

702

oc

n

7/ 1/29

18.3

34.3

8.18

4

llSa

705

oc

P

7/ 1/29

18.3

34.3

8.18

4

116

703

50

c

n

7/ 1/29

10.6

33.8

8.11

23

116

704

100

c

n

7/ 1/29

6.7

33.8

116

706

100

c

n

7/ 1/29

10.6

33.8

s.'ii

23

116

707

100

r

P

7/ 1/29

6.7

33.8

117

708

oc

n

7/ 3/29

15.9

34.3

8. '17

"3'

117

710

100

oc

n

7/ 3/29

8.8

34.1

7.98

84

117

711

oc

P

7/ 3/29

15.9

34.3

8.17

3

117

712

50

r

P

7/ 3/29

12.5

34.2

8.06

51

118

716

100

r

n

7/ 5/29

6.1

33.8

7.94

114

118

717

r

P

7/ 5/29

10.2

33.6

8.21

90

119

720

r

P

7/ 7/29

6.9

33.0

7.96

142

126

761

oc

n

7/21/29

11.2

32.6

8.09

76

127

767

oc

n

7/23/29

13.4

32.7

8.12

43

127

768

50

r

n

7/23/29

10.5

32.8

8.09

56

128

773

c

n

7/25/29

16.4

33.0

8.12

29

128

774

50

r

n

7/25/29

11.8

33.1

8.11

29

128

776

c

P

7/25/29

16.4

33.0

8.12

29

128

775

100

oc

n

7/25/29

10.2

33.2

8.06

46

128

777

50

r

P

7/25/29

11.8

33.1

8.11

29

128

778

100

r

P

7/25/29

10.2

33.2

8.06

46

128

779

oc

n

7/25/29

16.4

33.0

8.12

29

129

780

oc

n

7/27/29

16.3

33.1

8.13

25

130

781

a

n

9/ 4/29

16.2

33.4

8.34

26

130

782

50

c

n

9/ 4/29

11.7

33.4

8.26

83

130

783

100

c

n

9/ 4/29

8.8

33.7

8.06

176

130

784

c

p

9/ 4/29

16.2

33.4

8.34

26

130

785

50

c

P

9/ 4/29

11.7

33.4

8.26

83

130

786

100

r

P

9/ 4/29

8.8

33.7

8.06

176

130

787

c

n

9/ 4/29

16.2

33.4

8.34

36

131

788

r

n

9/ 6/29

19.3

33.4

8.34

131

789

100

oc

n

9/ 6/29

12.1

33.4

8.32

* . .

131

792

oc

p

9/ 6/29

19.3

33.4

8.34

...

131

793

r

P

9/ 6/29

19.3

33.4

8.34

...

131

794

r

P

9/ 6/29

19.3

33.4

8.34

• • .

131

795

r

P

9/ 6/29

19.3

33.4

8.34

132

797

oc

n

9/ 8/29

21.0

33.9

8.34

i'5

132

798

50

oc

n

9/ 8/29

17.6

33.9

8.33

19

132

799

100

oc

n

9/ 8/29

14.3

33.4

8.30

16

132

801

r

P

9/ 8/29

21.0

33.9

8.34

15

132

802

50

r

P

9/ 8/29

17.6

33.9

8.33

19

132

803

100

oc

P

9/ 8/29

14.3

33.4

8.30

16

143

881

oc

n

10/ 9/29

22.4

34.4

8.30

6

143

882

50

oc

n

10/ 9/29

19.0

34.2

8.34

6

143

883

100

oc

n

10/ 9/29

19.0

34.2

8.34

6

143

884

r

P

10/ 9/29

22.4

34.4

8.30

6

145

890

c

n

10/ 9/29

19.0

34.2

8.34

6

145

892

100

oc

n

10/ 9/29

16.0

34.1

8.31

6

145

893

oc

P

10/ 9/29

19.0

34.2

8.34

6

145

895

100

c

p

10/ 9/29

16.0

34.1

8.31

6

146

896

oc

n

10/15/29

22.4

34.9

8.37

6

146

897

50

oc

n

10/15/29

22.4

34.9

8.30

6

146

898

100

oc

n

10/15/29

19.7

34.3

8.26

7

146

901

50

r

P

10/15/29

22.4

34.9

8.30

6

147

905

100

r

n

10/15/29

19.2

35.0

8.29

5

78

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 23. Distributional and environmental records for Ceratium tripos subsp. semipulchellum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

(m)

abundance

ratus

(°C)

(o/oo)

mg/m^

1

1

oc

n

5/12/28

24.0

36.2

8.16

34

1

2

70

r

n

5/12/28

(22.4

36.4

8.15

36

la

4

c

n

5/18/28

(22.3

36.3

8.20

46)

la

5

c

n

5/18/28

22.3

36.3

8.20

46

lb

6

c

n

5/18/28

(22.3

36.3

8.20

46

lb

7

50

c

n

5/18/28

21.7

36.3

8.20

38

lb

8

100

oc

n

5/18/28

20.0

36.5

8.18

36)

2

12

50

00

n

5/18/28

20.5

36.4

8.21

46

14

101

50

r

n

8/ 9/28

15.0

35.1

8.18

16

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

16

104

50

r

P

8/13/28

24.4

36.4

8.23

8

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

17

111

r

n

8/15/28

26.2

36.6

8.29

9

17

112

50

r

n

8/15/28

21.9

36.6

8.28

12

17

113

100

r

n

8/15/28

19.3

36.5

8.23

9

20

125

r

P

8/22/28

26.0

36.6

8.37

5

20

128

oc

n

8/22/28

26.0

36.6

8.37

5

23

145

oc

P

8/29/28

27.2

35.9

8.25

4

23

148

oc

n

8/29/28

27.2

35.9

8.25

4

23

149

50

oc

n

8/29/28

20.9

36.0

8.14

13

23

150

100

r

n

9/29/28

16.6

36.0

8.18

75

24

151

oc

P

8/31/28

27.2

35.2

8.32

4

24

153

100

r

P

8/31/28

15.6

35.6

7.96

99

24

154

oc

n

8/31/28

27.2

35.2

8.32

4

24

156

100

r

n

8/31/28

15.6

35.6

7.96

99

25

161

50

oc

n

9/ 3/28

21.5

36.0

8.22

12

26

165

100

r

p

9/ 5/28

14.9

35.6

8.11

40

27

169

100

r

p

9/ 7/28

17.7

36.0

8.09

46

27

172

100

oc

n

9/ 7/28

17.7

36.0

8.09

46

28

174

50

r

p

9/11/28

26.7

36.3

8.26

4

28

177

50

r

n

9/11/28

26.7

36.3

8.26

4

29

180

50

r

p

9/13/28

27.2

36.2

8.29

3

29

182

r

n

9/13/28

27.6

36.2

8.31

3

30

186

50

oc

p

9/15/28

27.8

36.1

8.29

3

30

189

50

c

n

9/15/28

27.8

36.1

8.29

3

30

190

100

oc

n

9/15/28

24.1

36.4

8.10

20

31

191

oc

n 10/ 3/28

28.5

34.4

8.27

2

31

192

50

oc

n 10/ 3/28

28.2

35.4

8.23

2

32

195

50

r

n 10/ 5/28

27.2

36.0

8.24

2

33

198

50

r

n 10/ 8/28

28.2

36.2

8.24

4

33

199A

100

r

p 10/ 8/28

22.3

36.5

8.18

23

34

201

100

oc

n 10/ 9/28

20.5

36.6

8.16

16

34

202

50

oc

n 10/ 9/28

25.0

36.5

8.21

3

35

207

50

r

p 10/26/28

16.8

34.7

7.92

138

35a

208

oc

n 10/30/28

(26.5

31.6

8.23

16)

35-36

209

oc

n 10/26/28

(27.4

29.7

8.31

15

35-36

212

oc

n 10/30/28

(26.9

35.6

8.27

15)

36

213

r

n 10/30/28

26.5

31.6

8.23

16

36

215

100

r

n 10/30/28

14.4

34.9

7.85

149

36

216

oc

p 10/30/28

26.5

31.6

8.23

16

37

219

100

oc

n 11/ 1/28

15.1

34.9

7.82

153

40

229

r

n 11/ 8/28

22.2

33.7

8.21

24

40

231

100

r

n 11/ 8/28

13.9

35.0

7.85

159

40

232

r

p 11/ 8/28

22.2

33.7

8.21

24

40

233

50

r

p 11/ 8/28

15.3

34.9

7.87

161

40

234

100

r

p 11/ 8/28

13.9

35.0

7.85

159

41

235

r

n 11/10/28

20.4

34.2

8.11

32

41

237

100

r

n 11/10/28

14.5

35.0

7.91

152

41

238

r

p 11/10/28

20.4

34.2

8.11

32

41

239

100

r

p 11/10/28

14.5

35.0

7.91

152

41a

240

r

n 11/10/28

(20.4

34.2

8.11

32)

42

241

r

n 11/13/28

18.7

34.7

8.06

45

43

247

r

p 11/15/28

19.6

34.8

8.09

52

44

250

r

n 11/17/28

20.7

34.9

8.03

38

45

257

50

r

n 11/19/28

22.4

35.2

8.13

46

45

259

oc

p 11/19/28

22.4

35.3

8.12

38

46

261

oc

n 11/21/28

23.3

35.3

8.16

36

46

262

50

r

n 11/21/28

23.2

35.3

8.16

40

46

263

100

r

n 11/21/28

22.5

35.4

8.17

40

46

264

r

p 11/21/28

23.3

35.3

8.16

36

46

265

50

r

p 11/21/28

23.2

35.3

8.16

40

47

266

r

n 11/23/28

23.9

36.0

8.23

17

47

267

50

c

n 11/23/28

23.8

36.0

8.23

20

47

268

100

oc

n 11/23/28

22.7

36.2

8.23

20

47

269

oc

p 11/23/28

23.9

36.0

8.23

17

APPENDIX

79

Table 23. Distributional and environmental records for
Ceratium tripos subsp. semipulchelluin--Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

PH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

47

269

r

P

11/23/28

23.9

36.0

8.23

17

47

270

50

r

P

11/23/28

23.9

36.0

8.23

17

48

271

DC

n

11/25/28

23.6

36.4

8.23

13

48

272

50

00

n

11/25/28

23.6

36.4

8.24

16

48

273

100

r

n

11/25/28

22.7

36.3

8.26

16

48

274

r

P

11/25/28

23.6

36.4

8.23

13

49

277

r

n

11/27/28

23.4

36.2

8.27

13

49

281

50

r

P

11/27/28

22.6

36.1

8.26

13

56

330

100

oc

n

12/18/28

16.6

34.8

8.11

12

56

333

100

r

P

12/18/28

16.6

34.8

8.11

12

56-57

334

oc

n

12/20/28

(19.0

34.5

8.14

20)

57

335

oc

n

12/20/28

19.0

34.5

8.14

20

57

336

50

r

n

12/20/28

15.6

34.3

8.14

21

57

337

100

oc

n

12/20/28

14.3

34.4

8.10

40

57

338

r

F

12/20/28

19.0

34.5

8.14

20

57

340

100

r

P

12/20/28

14.3

34.4

8.10

40

58

344

r

P

12/22/28

17.0

34.0

8.12

20

59

348

r

n

12/24/28

16.3

34.0

8.10

38

59

350

100

r

n

12/24/28

11.4

34.1

8.03

72

59

351

r

P

12/24/28

16.3

34.0

8.10

38

60-61

361

r

n

12/24/28

(16.3

34.0

8.10

38)

61

364

50

a

n

12/28/28

14.0

34.0

8.10

60

61-62

369

oc

n

12/28/28

(16.9

34.0

8.05

46)
46

61-62

370

r

n

12/28/28

(16.9

34.0

8.05

62

372

50

c

n

12/30/28

16.2

34.3

8.10

28

62

373

100

oc

n

12/30/28

13.1

34.2

8.06

48

62

374

oc

P

12/30/28

19.2

34.2

8.12

32

62

376

100

r

P

12/30/28

13.1

3'i.2

8.06

48

62-63

378

r

n

1/ 1/29

(20.5

34.6

8.07

21)

63

379

oc

n

1/ 1/29

20.5

34.6

8.07

21

63

380

50

oc

n

1/ 1/29

17.0

34.6

8.08

25

63

382

r

P

1/ 1/29

20.5

34.6

8.07

21

63

383

50

oc

P

1/ 1/29

17.0

34.6

8.08

25

63-64

385

r

P

1/ 1/29

(20.5

34.6

8.07

21

)

63-64

387

r

n

1/ 1/29

20.5

34.6

8.07

21

63-64

388

oc

n

1/ 1/29

20.5

34.6

8.07

21

63-64

389

r

n

1/ 1/29

20.5

34.6

8.07

21

63-64

390

r

n

1/ 1/29

20.5

34.6

8.07

21

63-64

391

r

n

1/ 1/29

20.5

34.6

8.07

21

63-64

391

oc

n

1/ 1/29

(20.5

34.6

8.07

21

64

394

r

n

1/ 3/29

20.6

34.6

8.12

21

64

395

50

r

n

1/ 3/29

17.2

34.6

8.12

29

64

399

50

r

P

1/ 3/29

17.2

34.6

8.12

29

64-65

401

c

n

1/ 3/29

(20.6

34.6

8.12

21]

64-65

401

r

n

1/ 3/29

20.6

34.6

8.12

21

64-65

401

oc

n

1/ 3/29

20.6

34.6

8.12

21

64-65

402

oc

n

1/ 3/29

20.6

34.6

8.12

21

64-65

403

oc

n

1/ 3/29

20.6

34.6

8.12

21'

64-65

403

r

n

1/ 3/29

20.6

34.6

8.12

21'

64-65

404

r

n

1/ 3/29

(20.6

34.6

8.12

2l|

65

405

r

n

1/ 5/29

20.2

34.5

8.10

24

65

406

50

r

n

1/ 5/29

16.5

34.5

8.10

25

65

407

100

r

n

1/ 5/29

14.8

34.3

8.10

34

65

408

r

P

1/ 5/29

20.2

34.5

8.10

24

65

409

50

oc

P

1/ 5/29

16.5

34.5

8.10

25

65

410

100

r

P

1/ 5/29

14.8

34.3

8.10

34

66

411

r

n

1/ 7/29

19.4

34.7

8.10

29

66

412

100

r

n

1/ 7/29

17.8

34.9

8.12

21

66

413

r

P

1/ 7/29

19.4

34.7

8.10

29

66

414

50

r

P

1/ 7/29

17.8

34.8

8.10

29

66

415

100

r

P

1/ 7/29

17.8

34.9

8.12

21

67

416

c

n

1/ 8/29

19.3

34.9

8.11

21

67

418

100

oc

n

1/ 8/29

16.2

34.6

8.05

40

69

424

oc

n

1/12/29

21.1

35.2

8.12

62

69

'25

50

oc

n

1/12/29

17.4

35.1

7.99

151

69

426

100

r

n

1/12/29

14.6

34.8

7.86

198

70

431

r

n

1/13/29

21.2

35.1

8.05

103

70

432

50

r

n

1/13/29

15.4

35.0

7.88

178

70

434

r

P

1/13/29

21.2

35.1

8.05

103

71

438

oc

n

2/ 6/29

23.5

35.2

8.13

58

71

•139

r

n

2/ 6/29

23.5

35.2

8.13

58

72

444

50

r

n

2/ 8/29

18.7

35.4

8.12

60

72-73

449

r

n

2/10/29

(25.3

35.4

B.21

44)

73

451

50

oc

n

2/10/29

18.7

35.4

8.05

122

73

452

100

oc

n

2/10/29

14.7

35.0

7.80

178

80

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 23. Distributional and environmental records for
Ceratium tripos subsp. semlpulchellum --Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
CC)

Salinity
(o/oo)

pH

PO4
mg/m^

75

464

100

r

n

2/14/29

17.8

35.4

8.00

75

75

466

50

r

p

2/14/29

20.0

35.5

8.14

46

77

472

c

n

2/18/29

23.7

36.0

8.19

16

77

473

oc

P

2/18/29

23.7

36.0

8.19

16

77

474

50

oc

p

2/18/29

23.5

36.0

8.19

16

78

475

oc

n

2/20/29

24.6

36.0

8.17

32

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

478

oc

p

2/20/29

24.6

36.0

8.17

32

78

479

50

r

p

2/20/29

23.8

36.1

8.14

32

78

480

100

r

p

2/20/29

21.9

36.2

8.14

34

79

481

oc

n

2/22/29

25.2

36.0

8.17

34

79

482

50

oc

n

2/22/29

24.5

36.1

8.17

34

79

483

100

oc

n

2/22/29

21.8

36.2

8.13

45

79

484

oc

p

2/22/29

25.2

36.0

8.17

34

79

485

50

oc

p

2/22/29

24.5

36.1

8.17

34

80

486

oc

n

2/24/29

26.0

35.9

8.20

36

80

487

50

oc

n

2/24/29

25.9

36.0

8.19

29

80

488

oc

p

2/24/29

26.0

25.9

8.20

36

80

489

50

oc

p

2/24/29

25.9

36.0

8.19

29

80

490

100

oc

p

2/24/29

23.4

36.2

8.16

32

81

491

c

n

2/26/29

26.5

35.8

8.19

38

81

492

50

oc

n

2/26/29

26.4

35.9

8.19

38

81

493

oc

P

2/26/29

26.5

35.8

8.19

38

81

494

50

oc

p

2/26/29

26.4

35.9

8.19

38

81

495

100

r

P

2/26/29

23.6

36.2

8.18

36

82

497

50

oc

n

2/28/29

27.2

36.3

8.21

34

82

500

100

r

p

2/28/29

24.4

36.5

8.19

34

83

501

oc

n

3/ 2/29

27.5

36.3

8.24

29

83

502

50

c

n

3/ 2/29

27.4

36.5

8.24

25

83

503

r

p

3/ 2/29

27.5

36.3

8.24

29

84

505

r

n

3/ 4/29

27.8

36.2

8.23

24

84

506

50

oc

n

3/ 4/29

27.5

36.4

8.21

24

85

510

oc

n

3/ 6/29

27.9

36.2

8.22

40

85

511

50

oc

n

3/ 6/29

27.8

36.2

8.22

40

86

517

50

oc

n

3/ 9/29

27.4

36.2

8.22

17

88

526

oc

n

3/21/29

28.5

35.9

8.23

16

88

527

50

oc

n

3/21/29

28.4

35.9

8.25

13

89

528

oc

n

3/23/29

28.4

35.6

8.25

21

89

529

50

c

n

3/23/29

28.6

35.8

8.27

12

89

531

100

r

p

3/23/29

26.4

36.0

8.24

12

90

533

r

n

3/25/29

28.5

35.5

8.27

21

90

535

oc

p

3/25/29

28.5

35.5

8.27

21

91

540

r

n

3/27/29

28.7

35.1

8.30

21

91

541

50

c

n

3/27/29

28.5

35.2

8.30

24

92

546

50

c

n

3/29/29

28.4

35.4

8.29

28

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

550

c

n

3/31/29

28.7

34.7

8.30

28

93

551

50

c

n

3/31/29

28.5

34.8

8.30

28

93

552

oc

p

3/31/29

28.7

34.7

8.30

28

93

553

100

r

p

3/31/29

27.6

35.8

8.27

29

94

558

50

oc

n

4/22/29

29.3

34.7

8.25

14

94

559

100

oc

n

4/22/29

28.5

35.6

8.21

25

95

563

50

c

n

4/24/29

29.3

34.9

8.24

16

95

564

100

c

n

4/24/29

28.5

35.4

8.22

21

95

566

50

r

P

4/24/29

29.3

34.9

8.24

16

95

587

100

r

P

4/24/29

28.5

35.4

8.22

21

96

568

c

n

4/26/29

29.3

35.3

8.23

12

95

568A

oc

n

4/24/29

(29.4

34.7

8.26

14)

96

569

50

oc

n

4/26/29

29.2

35.3

8.23

12

96

570

100

oc

n

4/26/29

28.2

35.7

8.19

25

96

571

r

P

4/26/29

29.3

35.3

8.23

12

96

573

100

r

P

4/26/29

28.2

35.7

8.19

25

97

575

50

c

n

4/28/29

28.0

35.4

8.16

21

97

576

100

oc

n

4/28/29

27.6

35.6

8.15

25

97

578

50

r

P

4/28/29

28.0

35.4

8.16

21

97

579

100

r

p

4/28/29

27.6

35.6

8.15

25

98

581

oc

n

4/30/29

27.0

35.3

8.16

24

98

582

50

oc

n

4/30/29

26.9

35.3

8.16

28

98

584

oc

p

4/30/29

27.0

35.3

8.16

24

98

585

50

r

p

4/30/29

26.9

35.3

8.16

28

98

588

100

r

p

4/30/29

26.7

35.4

8.14

32

99

589

c

n

5/ 2/29

27.9

34.9

8.21

12

99

590

50

c

n

5/ 2/29

27.8

34.9

8.22

12

99

591

100

c

n

5/ 2/29

27.8

35.0

8.22

17

APPENDIX

81

Table 23. Distributional and environmental records for
Ceratlum tripos subsp. semipulcheUuin--Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

99

592

oc

P

5/ 2/29

27.9

34.9

8.21

12

99

592

r

P

5/ 2/29

27.9

34.9

8.21

12

99

593

50

r

P

5/ 2/29

27.8

34.9

8.22

12

99

594

100

r

P

5/ 2/29

27.8

35.0

8.22

17

99

595

c

n

5/ 2/29

27.9

34.9

8.21

12

100

596

50

c

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

c

n

5/ 4/29

27.6

34.7

8.22

12

100

598

oc

P

5/ 4/29

27.7

34.7

8.21

10

100

599

50

oc

P

5/ 4/29

27.6

34.7

8.21

10

100

602

100

oc

P

5/ 4/29

27.6

34.7

8.22

12

101

603

c

n

5/ 7/29

26.3

34.7

8.24

8

101

606

r

P

5/ 7/29

26.3

34.7

8.24

8

101

607

50

r

P

5/ 7/29

26.2

34.7

8.24

8

102

609

c

n

5/ 9/29

25.8

35.0

8.24

8

102

610

50

c

n

5/ 9/29

25.8

35.0

8.24

8

102

611

100

c

n

5/ 9/29

25.6

35.0

8.23

8

102

612

r

P

5/ 9/29

25.8

35.0

8.24

8

102

613

50

oc

P

5/ 9/29

25.8

35.0

8.24

8

103

615

c

n

5/11/29

26.0

35.0

8.25

5

103

616

50

c

n

5/11/29

25.8

35.2

8.25

5

103

617

100

c

n

5/11/29

24.8

35.2

8.25

5

103

618

r

P

5/11/29

26.0

35.0

8.25

5

103

620

100

r

P

5/11/29

24.8

35.2

8.25

5

104.

621

c

n

5/13/29

26.1

35.2

8.24

7

104

622

50

c

n

5/13/29

25.8

35.2

8.24

7

104

623

100

c

n

5/13/29

25.3

35.3

8.21

7

104

625

SO

r

P

5/13/29

25.8

35.2

8.24

7

104

626

100

r

P

5/13/29

25.3

35.3

8.21

7

105

628

50

c

n

5/15/29

26.8

34.9

8.23

5

105

629

100

oc

n

5/15/29

25.2

35.1

8.23

5

105

630

r

P

5/15/29

26.9

34.9

8.23

5

106

633

oc

n

5/17/29

27.2

35.0

8.23

5

106

634

50

c

n

5/17/29

27.0

35.0

8.23

5

106

635

100

c

n

5/17/29

25.6

35.1

8.23

5

107

639

c

n

5/19/29

28.0

34.4

8.23

5

107

640

SO

c

n

5/19/29

27.9

34.4

8.23

4

107

641

100

c

n

5/19/29

26.8

34.9

8.23

4

108

646

oc

n

5/27/29

28.4

35.0

8.25

4

108

647

SO

c

n

5/27/29

26.8

35.0

8.24

4

108

648

100

c

n

5/27/29

25.2

35.0

8.23

4

109

656

100

oc

n

5/29/29

19.4

34.8

8.18

5

109

658

so

r

p

5/29/29

23.1

35.0

8.22

3

109

659

100

r

P

5/29/29

19.4

34.8

8.18

5

110

661

r

n

5/31/29

23.9

34.7

8.18

5

110

661

oc

P

5/31/29

23.9

34.7

8.18

5

111

664

a

n

6/ 3/29

20.1

34.5

8.18

5

111

' 669

50

oc

n

6/ 3/29

19.4

34.6

8.17

5

111

670

100

r

n

6/ 3/29

18.2

34.7

8.13

13

111

671

oc

p

6/ 3/29

20.1

34.5

8.18

5

111

672

50

oc

p

6/ 3/29

19.4

34.6

8.17

5

111

674

c

n

6/ 3/29

20.1

34.5

8.18

5

112

675

SO

c

n

6/ 5/29

21.7

34.6

8.23

7

112

676

100

oc

n

6/ 5/29

19.8

34.7

8.20

8

112

677

oc

P

6/ 5/29

23.2

34.8

8.22

7

113

680

c

n

6/25/29

24.2

34.5

8.25

5

lis

681

50

c

n

6/25/29

23.8

34.6

8.25

5

113

682

100

c

n

6/25/29

21.5

34.7

8.23

8

lis

683

oc

P

6/25/29

24.2

34.5

8.25

5

114

690

50

oc

P

6/27/29

16.2

34.6

8.04

63

115

692

oc

n

6/29/29

20.6

34.6

8.19

115

693

50

r

n

6/29/29

17.5

34.6

8.12

17

115

694

100

r

n

6/29/29

15.6

34.6

8.08

27

115

695-6

oc

P

6/29/29

20.6

34.6

8.19

117

709

50

oc

n

7/ 3/29

12.5

34.2

8.06

51

133

804

oc

n

9/10/29

22.7

34.7

8.47

133

80S

50

oc

n

9/10/29

20.8

34.7

8.37

133

806

100

r

n

9/10/29

18.4

34.8

8.31

134

808

c

n

9/12/29

22.9

34.7

8.34

6

134

809

SO

oc

n

9/12/29

19.8

34.6

8.34

6

134

810

100

oc

n

9/12/29

18.1

34.6

8.34

6

133

812

oc

P

9/10/29

22.7

34.7

8.47

7

133

814

100

oc

P

9/10/29

18.4

34.8

8.31

7

134

815

oc

P

9/12/29

22.9

34.7

8.34

6

134

817

100

oc

p

9/12/29

18.1

34.6

8.34

6

135

822

100

oc

n

9/14/29

18.7

34.8

8.34

5

82

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 23. Distributional and environmental records for
Ceratium tripos subsp. semipulchellum- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

135

824

r

p

9/14/29

23.8

35.1

8.37

7

135

826

100

r

P

9/14/29

18.7

34.8

8.34

5

136

827

oc

n

9/16/29

24.6

35.4

8.37

3

136

828

50

oc

n

9/16/29

21.4

35.1

8.39

3

136

832

r

P

9/16/29

24.6

35.4

8.37

3

136

829

100

oc

n

9/16/29

18.6

35.0

8.39

3

136

833

50

r

P

9/16/29

21.4

35.1

8.39

3

136

834

100

r

p

9/16/29

18.6

35.0

8.39

3

137

836

c

n

9/18/29

25.5

35.0

8.39

4

137

837

c

n

9/18/29

25.5

35.0

8.39

4

137

840

r

P

9/18/29

25.5

35.0

8.39

4

137

841

oc

P

9/18/29

25.5

35.0

8.39

4

138

843

oc

n

9/20/29

26.1

34.8

8.35

5

138

845

100

c

n

9/20/29

22.2

34.8

8.31

3

138

848

50

r

P

9/20/29

26.1

. 34.8

8.35

5

139

849

r

n

9/22/29

26.7

34.8

8.34

6

139

850

50

c

n

9/22/29

25.8

34.9

8.31

6

139

851

100

oc

n

9/22/29

22.4

35.2

8.28

6

139

854

50

r

P

9/22/29

25.8

34.9

8.31

6

139

855

100

r

p

9/22/29

22.4

35.2

8.28

6

140

856

c

n

10/ 3/29

26.9

35.0

8.42

7

140

857

50

oc

n

10/ 3/29

26.9

35.0

8.39

7

140

858

100

oc

n

10/ 3/29

25.5

35.0

8.34

7

140

860

r

P

10/ 3/29

26.9

35.0

8.42

7

140

861

50

r

p

10/ 3/29

26.9

35.0

8.39

7

140

863

100

r

P

10/ 3/29

25.5

35.0

8.34

7

141

864

c

n

10/ 5/29

25.9

35.2

8.34

5

141

865

50

c

n

10/ 5/29

24.8

35.3

8.34

5

141

871

100

r

P

10/ 5/29

20.0

35.0

8.33

5

142

873

50

oc

n

10/ 5/29

20.0

35.0

8.33

5

142

874

100

c

n

10/ 5/29

16.6

34.4

8.27

7

142

876

oc

P

10/ 5/29

24.1

34.8

8.33

5

142

878

100

r

P

10/ 5/29

16.6

34.4

8.27

7

142

879

100

r

P

10/ 5/29

16.6

34.4

8.27

7

143

881

c

n

10/ 9/29

22.4

34.4

8.30

6

143

882

50

oc

n

10/ 9/29

19.0

34.2

8.34

6

14S

883

100

oc

n

10/ 9/29

13.8

34.1

8.30

10

143

884

oc

P

10/ 9/29

22.4

34.4

8.30

6

144

886

c

n

10/11/29

23.3

35.0

8.37

6

144

887

oc

P

10/11/29

23.3

35.0

8.37

6

144

889

100

r

P

10/11/29

16.6

34.5

8.37

6

145

891

50

c

n

10/11/29

18.7

34.3

8.34

6

145

894

50

r

P

10/11/29

18.7

34.3

8.34

6

146

896

oc

n

10/15/29

22.4

34.9

8.37

6

146

897

50

oc

n

10/15/29

22.4

34.9

8.30

6

146

898

100

oc

n

10/15/29

19.7

34.3

8.26

7

146

901

50

r

P

10/15/29

22.4

34.9

8.30

6

147

903

oc

n

10/17/29

23.3

35.3

8.26

8

147

904

50

r

n

10/17/29

23.1

35.3

8.29

5

147

907

oc

p

10/17/29

23.3

35.3

8.26

8

149

920

oc

n

10/21/29

23.5

35.0

8.34

6

149

921

50

oc

n

10/21/29

23.3

35.0

8.37

6

149

922

100

oc

n

10/21/29

20.3

34.9

8.38

6

149

928

50

r

p

10/21/29

23.3

35.0

8.37

6

150

929

oc

n

10/23/29

25.6

34.7

8.39

7

150

930

50

r

n

10/23/29

27.8

34.8

8.35

10

150

931

100

r

n

10/28/29

19.6

34.6

8.32

11

161

938

50

c

n

10/26/29

18.3

34.4

• ••

151

939

100

oc

n

10/26/29

12.5

34.6

...

151

942

50

oc

p

10/26/29

18.3

34.4

.>•

152

945

50

oc

n

10/27/29

14.2

34.5

7.87

53

152

949

50

r

p

10/27/29

14.2

34.5

7.87

53

152

950

100

r

P

10/27/29

11.4

34.7

7.76

75

153

951

oc

n

10/27/29

28.1

34.2

8.47

153

952

50

c

n

10/29/29

28.1

34.4

8.39

153

953

100

oc

n

10/29/29

20.5

34.7

8.28

31

153

955

r

P

10/29/29

28.1

34.2

8.47

153

956

50

oc

P

10/29/29

28.1

34.4

8.39

153

957

100

oc

P

10/29/29

20. S

34.7

8.28

31

154

958

oc

n

10/31/29

28.3

34.2

8.39

154

959

50

oc

n

10/31/29

28.2

34.2

8.40

154

960

100

oc

n

10/31/29

25.3

34.8

7.93

21

155

965

c

n

11/ 2/29

27.8

34.9

8.29

29

155

966

50

c

n

11/ 2/29

27.7

34.9

8.30

30

155

967

100

oc

n

11/ 2/29

27.2

35.0

■8.30

35

APPENDIX

83

Table 23. Distributional and environmental records for
Ceratliun tripos subsp. semlpulchellum — Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

155

973

50

r

P

11/ 2/29

27.7

34.9

8.30

30

156

972

oc

n

11/ 4/29

27.6

35.0

8.34

28

156

972

r

n

11/ 4/29

27.6

35.0

8.34

28

156

974

100

oc

n

11/ 4/29

26.4

35.1

8.30

48

157

978

oc

n

11/ 6/29

27.1

35.3

8.27

47

157

979

50

c

n

11/ 6/29

27.1

35.2

8.32

60

167

980

100

r

n

11/ 6/29

26.8

35.5

8.30

64

158

983

c

n

11/ 8/29

28.2

35.6

8.34

36

158

984

50

c-

n

11/ 8/29

28.2

35.6

8.39

50

158

985

100

c

n

11/ 8/29

27.6

35.9

8.39

48

159

990

c

n

11/11/29

28.6

35.7

8.37

15

159

991

50

c

n

11/11/29

28.5

35.7

8.39

15

159

992

100

r

n

11/11/29

28.0

35.7

8.37

23

160

1000

r

n

11/13/29

28.6

35.6

8.37

12

160

1002

50

oc

n

11/13/29

28.6

35.6

8.39

15

160

1003

100

r

n

11/13/29

28.5

35.7

8.44

16

Table 24. Distributional and environmental records for Ceratlum pulchellum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/mS

14

93

oc

P

8/ 9/28

21.2

35.2

8.18

11

15

101

50

r

n

8/11/28

19.8

36.5

8.21

8

15

102

100

r

n

8/11/28

18.4

36.4

8.20

19

16

103

r

P

8/13/28

25.9

36.2

8.24

8

17

109

r

p

8/15/28

26.2

36.6

8.29

9

18

114

oc

P

8/17/28

27.0

37.0

8.23

5

19

123

50

r

n

8/20/28

25.2

37.1

8.27

5

20

125

r

p

8/22/28

26.0

36.6

8.37

5

SO

126

SO

r

p

8/22/28

25.8

36.6

8.26

3

20

127

100

r

p

8/22/28

22.6

36.7

8.19

5

20-21

131

r

n

8/24/28

(26.6

36.3

8.32

4)

21

134

100

r

p

8/24/28

21.0

36.8

8.20

7

21

136

50

r

n

8/24/28

24.4

36.2

8.26

4

21

137

100

r

n

8/24/28

21.0

36.8

8.20

7

22

144

100

r ■

n

8/27/28

17.5

36.1

7.99

123

23

145

r

p

8/29/28

27.2

35.9

8.25

4

48

272

50

r

n

11/25/28

23.6

36.4

8.24

16

48

274

oc

p

11/25/28

23.6

36.4

8.24

16

48

275

50

oc

p

11/25/28

23.6

36.4

8.24

16

49

260

r

p

11/27/28

23.4

36.2

8.27

13

49a

283

p

n

11/29/28

(23.2

36.0

8.23

13)

50

285

50

p

n

11/29/28

22.0

35.9

8.23

13

50

287

oc

p

11/29/28

23.2

36.0

8.23

13

50-51

290

oc

n

12/ 1/28

(22.8

35.6

8.22

16}

50-51

291

oc

n

12/ 1/28

22.8

35.6

8.22

16

50-51

292

oc

n

12/ 1/28

122.8

35.6

8.22

16)

51

294

50

oc

n

12/ 1/28

20.5

35.6

8.22

17

51

296

r

p

12/ 1/28

22.8

35.6

8.22

16

51

297

50

p

p

12/ 1/28

20.5

35.6

8.22

17

52

300

50

r

n

12/ 3/28

20.2

35.6

8.20

8

52

301

100

p

n

12/ 3/28

18.2

35.2

8.17

8

52

302

oc

p

12/ 3/28

22.5

35.4

8.21

8

53

304

50

p

n

12/ 5/28

21.2

35.8

8.20

IS

53

305

100

p

n

12/ 5/28

19.9

35.6

8.19

13

53

306

r

p

12/ 5/28

22.6

35.7

8.22

13

53-54

310

r

n

12/14/28

(83.0

35.6

8.22

11

53-54

311

r

n

12/14/28

23.0

35.6

8.22

11

53-54

313

r

n

12/14/38

123.0

35.6

8.22

11

54

320

50

r

n

12/14/28

19.8

35.4

8.18

17

54

321

100

p

n

12/14/28

18.7

35.4

8.16

20

54

322

oc

p

12/14/28

23.2

35.5

8.22

9

55

323

p

n

12/16/28

20.4

34.9

8.19

12

55

324

50

p

n

12/16/28

18.7

35.0

8.18

12

55

325

100

p

n

12/16/28

16.7

34.9

7.17

12

56

328

oc

n

12/18/28

20.8

34.9

8.13

9

56

330

100

oc

n

12/18/28

16.6

34.8

8.11

12

56

331

r

p

12/18/28

20.8

34.9

8.13

9

56-57

334

c

n

12/20/28

(19.0

34.5

8.14

20

61-62

369

c

n

12/28/28

16.9

34.0

8.05

46)

61-62

370

oc

n

12/28/28

1I6.9

34.0

8.05

46

84

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 24. Distributional and environmental records for Ceratium piilchellum — Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PC

(m)

abundance

ratus

(°C)

(o/oo)

mg/m^

62

372

50

oc

n

12/30/28

16.2

34.3

8.10

28

62

373

100

DC

n

12/30/28

13.1

34.2

8.06

48

62

374

oc

P

12/30/28

19.2

34.2

8.12

32

62

375

50

oc

P

12/30/28

16.2

34.3

8.10

28

62-63

377

oc

n

1/ 1/29

(20.5
(20.5

34.6

8.07

21)
21

62-63

378

r

n

1/ 1/29

34.6

8.07

63

379

oc

n

1/ 1/29

20.5

34.6

8.07

21

63

381

100

r

n

1/ 1/29

15.6

34.6

8.08

24

63

382

oc

P

1/ 1/29

20.5

34.6

8.07

21

63

384

100

r

P

1/ 1/29

15.6

34.6

8.08

24

63-64

385

r

P

1/ 1/29

(20.5

34.6

8.07

21)

63-64

386

c

n

1/ 1/29

120.5

34.6

8.07

21i

63-64

387

c

n

1/ 1/29

120.5

34.6

8.07

21;

63-64

388

oc

n

1/ 1/29

20.5

34.6

8.07

21)

63

389

oc

n

1/ 1/29

(20.5

34.6

8.07

21)

63-64

390

oc

n

1/ 1/29

20.5

34.6

8.07

21

63-64

391

c

n

1/ 1/29

(20.5

34.6

8.07

21

63-64

392

oc

n

1/ 1/29

20.5

34.6

8.07

21

63-64

393

oc

n

1/ 1/29

120.5

34.6

8.07

21)

64

394

oc

n

1/ 3/29

20.6

34.6

8.12

21

64

395

50

r

n

1/ 3/29

17.2

34.6

8.12

29

64

396

100

oc

n

1/ 3/29

15.8

34.5

8.10

32

64

397

100

r

n

1/ 3/29

15.8

34.5

8.10

32

64

398

c

P

1/ 3/29

20.6

34.6

8.12

21

64

399

50

oc

P

1/ 3/29

17.2

34.6

8.12

29

64

400

100

r

P

1/ 3/29

15.8

34.5

8.10

32

64-65

401

oc

n

1/ 3/29

(15.8

34.5

8.10

32)

64-65

402

oc

n

1/ 3/29

(15.8

34.5

8.10

32)

64-65

403

oc

n

1/ 3/29

(15.8

34.5

8.10

32)

68

422

r

P

1/10/29

19.2

35.1

8.14

29

84

506

50

oc

n

3/ 4/29

27.5

36.4

8.21

24

84

507

r

p

3/ 4/29

27.8

36.2

8.23

24

86

516

r

n

3/ 9/29

28.3

36.2

8.29

20

86

518

oc

p

3/ 9/29

28.3

36.2

8.29

20

86

519

50

r

P

3/ 9/29

27.4

36.2

8.29

17

87

523

oc

p

3/ 9/29

28.3

36.2

8.29

20

87

525

100

r

p

3/11/29

23.9

36.0

8.23

20

89

528

r

n

3/23/29

28.4

35.6

8.25

21

89

529

50

oc

n

3/23/29

28.6

35.8

8.27

12

89

530

r

P

3/23/29

28.4

35.6

8.25

21

90

535

r

p

3/23/29

28.5

35.5

8.27

21

90

536

50

r

P

3/23/29

28.6

35.6

8.26

21

91

542

r

p

3/23/29

28.7

35.1

8.30

21

91

543

50

r

p

3/27/29

28.5

35.2

8.30

24

91

544

100

r

p

3/27/29

25.8

36.0

8.25

30

92

545

00

r

n

3/29/29

28.5

35.3

8.29

28

93

552

oc

p

3/29/29

28.7

34.7

8.29

28

94

558

50

oc

n

4/22/29

29.3

34.7

8.25

14

95

565

r

p

4/22/29

29.4

34.7

8.26

14

95

568A

r

n

4/24/29

(29.4

34.7

8.26

14)

96

571

r

p

4/26/29

29.3

35.3

8.23

12

101

605

100

r

n

5/ 7/29

25.2

35.1

8.23

8

101

606

r

P

5/ 7/29

26.3

34.7

8.24

8

102

610

50

c

n

5/ 2/29

25.8

35.0

8.24

8

102

612

r

P

5/ 9/29

25.8

35.0

8.24

8

102

614

100

r

P

5/ 9/29

25.6

35.0

8.23

8

104

624

r

P

5/13/29

26.1

35.2

8.24

7

105

627

50

c

n

5/15/29

26.8

34.9

8.23

5

106

635

100

c

n

5/17/29

25.6

35.1

8.23

5

106

636

r

P

5/17/29

27.2

35.0

8.23

5

107

642

oc

P

5/17/29

28.0

34.4

8.23

5

107

643

50

r

p

5/17/29

27.9

34.4

8.23

107

645

50

r

P

5/17/29

27.9

34.4

8.23

108

649

r

P

5/17/29

28.4

35.0

8.25

108

651

100

r

P

5/17/29

25.2

35.0

8.23

112

677

oc

P

5/17/29

23.2

34.6

8.22

135

820

r

n

9/14/29

23.8

35.1

8.37

135

822

100

oc

n

9/14/29

18.7

34.8

8.34

5

135

824

r

P

9/14/29

23.8

35.1

8.37

7

135

825

50

r

P

9/14/29

21.5

35.0

8.37

5

135

826

100

r

p

9/14/29

18.7

34.8

8.34

5

136

827

oc

n

9/16/29

24.6

35.4

8.37

3

136

832

r

p

9/16/29

24.6

35.4

8.37

3

136

833

50

r

P

9/16/29

21.4

35.1

8.39

3

136

835

100

r

P

9/16/29

21.4

35.1

8.39

3

137

838

100

r

n

9/16/29

21.5

35.1

8.30

5

APPENDIX

85

Table 24. Distributional and environmental records for Ceratlum pulchellum- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

138

843

r

n

9/20/29

26.1

34.8

8.35

5

138

844

50

r

P

9/20/20

26.1

34.8

8.35

5

138

847

r

P

9/20/29

26.1

34.8

8.35

5

141

868

r

P

9/20/29

26.1

34.8

8.35

5

141

870

50

r

P

9/20/29

26.1

34.8

8.35

5

142

872

r

n

10/ 7/29

24.1

34.8

8.33

5

142

873

50

r

n

10/ 7/29

21.8

34.8

8.30

5

142

876

r

P

10/ 7/29

24.1

34.8

8.33

5

142

878

100

r

P

10/ 7/29

16.6

34.4

8.27

7

143

881

c

n

10/ 9/29

22.4

34.4

8.30

6

143

883

100

r

n

10/ 9/29

13.8

34.1

8.30

10

144

886

r

n

10/11/29

23.3

35.0

8.37

6

144

887

DC

P

10/11/29

23.3

35.0

8.37

6

144

888

50

r

P

10/11/29

21.1

34.7

8.33

6

145

890

c

n

10/13/29

22.3

34.6

8.29

6

145

891

50

c

n

10/13/29

18.7

34.3

8.34

6

145

892

100

oc

n

10/13/29

16.0

34.1

8.31

6

145

893

oc

P

10/13/29

22.3

34.6

8.29

6

145

894

50

r

P

10/13/29

18.7

34.3

8.34

6

145

895

100

c

P

10/13/29

16.0

34.1

8.31

6

146

897

50

oc

n

10/15/29

22.4

34.9

8.37

6

146

900

r

P

10/15/29

22.4

34.9

8.37

6

146

901

50

r

P

10/15/29

22.4

34.9

8.30

6

148

910

oc

n

10/19/29

23.4

35.2

148

911

50

r

n

10/19/29

23.0

35.1

>■>

148

917

r

P

10/19/29

23.4

35.2

149

920

r

n

10/21/29

23.5

35.0

8.34

6

149

927

r

P

10/21/29

23.5

35.0

8.34

6

149

928

50

r

P

10/21/29

23.5

35.0

8.34

6

151

937

r

n

10/26/29

26.0

34.0

151

941

r

P

10/26/29

26.0

34.0

152

946

100

oc

n

10/26/29

11.4

34.7

7.'76

75

159

991

50

r

n

10/26/29

11.4

34.7

7.76

75

Table 25. Distributional and environmental records for Ceratlum breve

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

(°C)

(0/00)

mg/m^

1

1

r

n

5/12/28

24.0

36.2

8.16

34

2

12

50

r

n

5/18/28

20.5

36.4

8.21

46

16

103

r

P

8/13/28

25.9

36.2

8.24

8

16

106

oc

n

8/13/28

25.9

36.2

8.24

8

16

108

100

r

n

8/13/28

19.9

36.5

8.17

13

20-21

131

c

n

8/23/28

(26.6

36.3

8.32

4)

21

132

c

p

8/24/28

26.6

36.3

8.32

4

21

134

100

c

p

8/24/28

21.0

36.8

8.20

7

21

135

oc

n

8/24/28

26.6

36.3

8.32

4

21

137

100

oc

n

8/24/28

21.0

36.8

8.20

7

22

139

oc

p

8/27/28

26.7

36.0

8.26

8

22

144

100

r

n

8/27/28

17.5

36.1

7.99

123

23

145

c

p

8/29/28

27.2

35.9

8.25

4

23

149

50

r

n

8/29/28

20.9

36.0

8.14

13

23

150

100

r

n

8/29/28

16.6

36.0

8.18

75

24

151

c

p

8/31/28

27.2

35.2

8.32

4

24

152

50

r

p

8/31/28

23.1

36.0

8.14

8

24

153

100

r

p

8/31/28

15.6

35.6

7.96

99

24

154

r

n

8/31/28

27.2

35.2

8.32

4

24

155

50

r

n

8/31/28

23.1

36.0

8.14

8

24

156

100

r

n

8/31/28

15.6

35.6

7.96

99

25

157

c

p

9/ 3/28

27.5

35.6

8.31

5

25

160

r

n

9/ 3/28

27.5

35.6

8.31

5

25

161

50

r

n

9/ 3/28

21.5

36.0

8.22

12

25

162

100

r

n

9/ 3/28

14.6

35.7

7.93

121

26

163

oc

p

9/ 5/28

27.6

36.0

8.30

5

26

164

50

r

p

9/ 5/28

24.1

36.1

8.21

5

27

167

r

p

9/ 7/28

27.5

36.3

8.31

4

27

172

100

r

n

9/ 7/28

17.7

36.0

8.09

46

28

175

100

r

p

9/11/28

22.8

36.6

8.22

7

28

177

50

r

n

9/11/28

26.7

36.3

8.26

4

29

179

oc

p

9/13/28

27.6

36.2

8.31

3

29

182

oc

n

9/13/28

27.6

36.2

8.31

3

29

183

50

r

n

9/13/28

27.2

36.2

8.29

3

86

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 25. Distributional and environmental records for Ceratium breve- -Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

30

188

r

n

9/15/28

28.0

36.1

8.30

2

30

189

50

c

n

9/15/28

27.8

36.1

8.29

3

30

190

100

oc

n

9/15/28

24.1

36.4

8.10

20

31

191

r

n

10/ 3/28

28.5

34.4

8.27

2

31

193

100

r

n

10/ 3/28

23.4

36.5

8.19

28

32

194

oc

n

10/ 5/28

28.0

36.0

8.23

2

32

194A

r

p

10/ 5/28

28.0

36.0

8.23

2

32

195

50

r

n

10/ 5/28

27.2

36.0

8.24

2

33

197A

r

p

10/ 8/28

28.5

35.6

8.23

4

34

200

r

n

10/ 9/28

28.5

35.9

8.28

2

35

205

100

r

n

10/26/28

14.4

34.9

7.88

189

35

206

c

p

10/26/28

27.4

29.7

8.31

15

35

207

50

r

P

10/26/28

16.8

34.7

7.92

138

35a

208

c

n

10/26/28

(27.4

29.7

8.31

15)

35-36

209

c

n

10/27/28

27.4

29.7

8.31

15'

35-36

211

oc

n

10/28/28

27.0

30.7

8.27

16

35-36

212

c

n

10/28/28

27.0

30.7

8.27

16)

36

213

oc

n

10/30/28

26.5

31.6

8.23

16

36

216

oc

P

10/30/28

26.5

31.6

8.23

16

37

217

oc

n

11/ 1/28

27.1

31.7

8.28

15

37

218

50

oc

n

11/ 1/28

18.8

34.5

8.00

121

37

219

100

oc

n

11/ 1/28

15.1

34.9

7.82

153

37

220

oc

P

11/ 1/28

27.1

31.7

8.28

15

39

226

50

r

n

11/ 6/28

16.3

34.6

7.92

48

41

235

r

n

11/10/28

20.4

34.2

8.11

32

41a

240

oc

n

11/12/28

(18.7

34.7

8.06

45)

42

241

r

n

11/13/28

18.7

34.7

8.06

45

42

242

50

r

n

11/13/28

17.2

34.9

7.99

68

42

243

100

r

n

11/13/28

13.8

35.0

7.91

150

43

245

50

r

n

11/15/28

17.0

34.9

7.93

80

43

247

r

p

11/15/28

19.6

34.8

8.09

52

44

250

c

n

11/17/28

20.7

34.9

8.03

38

44

251

50

oc

n

11/17/28

20.4

34.9

8.04

34

44

252

100

oc

n

11/17/28

13.8

35.0

7.85

70

44

253

c

p

11/17/28

20.7

34.9

8.03

38

44

254

50

r

P

11/17/28

20.4

34.9

8.04

34

44

255

100

r

p

11/17/28

13.8

35.0

7.85

70

45

256

oc

n

11/19/28

22.4

35.3

8.12

38

45

257

50

oc

n

11/19/28

22.4

35.2

8.13

46

45

258

100

oc

n

11/19/28

18.6

35.1

8.00

50

45

259

oc

P

11/19/28

22.4

35.3

8.12

38

45

260

50

r

P

11/19/28

22.4

35.2

8.13

46

46

261

oc

n

11/21/28

23.3

35.3

8.16

36

46

262

50

oc

n

11/21/28

23.2

35.3

8.16

40

46

263

100

r

n

11/21/28

22.5

35.4

8.17

40

46

264

r

p

11/21/28

23.3

35.3

8.16

36

47

266

oc

n

11/23/28

23.9

36.0

8.23

17

47

267

50

oc

n

11/23/28

23.8

36.0

8.23

20

47

268

100

oc

n

11/23/28

22.7

36.2

8.23

20

47

269

oc

p

11/23/28

23.9

36.0

8.23

17

47

270

50

r

p

11/23/28

23.8

36.0

8.23

20

48

271

oc

n

11/25/28

23.6

36.4

8.23

13

48

272

SO

oc

n

11/25/28

23.6

36.4

8.24

16

48

273

100

r

n

11/25/28

22.7

36.3

8.26

16

49

277

r

n

11/27/28

23.4

36.2

8.27

13

49

279

100

r

n

11/27/28

21.8

35.9

8.26

13

49

280

r

p

11/27/28

23.4

36.2

8.27

13

SO

284

r

n

11/29/28

23.2

36.0

8.23

13

50

288

50

r

p

11/29/28

22.0

35.9

8.23

13

70

433

100

r

n

1/13/29

12.6

34.8

7.68

233

70

434

r

p

1/13/29

21.2

35.1

8.05

103

77

473

r

p

2/18/29

23.7

36.0

8.19

16

77

474

50

oc

p

2/18/29

23.5

36.0

8.19

16

78

475

oc

n

2/20/29

24.6

36.0

8.17

32

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

478

oc

p

2/20/29

24.6

36.0

8.17

32

78

479

SO

r

p

2/20/29

23.8

36.1

8.14

32

78

480

100

r

p

2/20/29

21.9

36.2

8.14

34

79

481

oc

n

2/22/29

25.2

36.0

8.17

34

79

482

50

oc

n

2/22/29

24.5

36.1

8.17

34

79

483

100

oc

n

2/22/29

21.8

36.2

8.13

45

79

484

oc

p

2/22/29

2S.2

36.0

8.17

34

80

486

oc

n

2/24/29

26.0

35.9

8.20

36

80

488

oc

P

2/24/29

26.0

35.9

8.20

36

80

489

50

oc

P

2/24/29

25.9

36.0

8.19

29

APPENDIX

87

Table 25. Distributional and environmental records for Ceratium breve- -Continued

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

PO4

(m)

abundance

ratus

CC)

(o/oo)

mg/m3

80

490

100

DC

P

2/24/29

23.4

36.2

8.16

32

81

491

DC

n

2/26/29

26.5

35.8

8.19

38

81

492

SO

oc

n

2/26/29

26.4

35.9

8.19

38

81

493

r

P

2/26/29

26.5

35.8

8.19

38

81

494

50

r

P

2/26/29

26.4

35.9

8.19

38

82

496

oc

n

2/28/29

27.2

36.3

8.21

34

82

497

SO

oc

n

2/28/29

27.2

36.3

8.21

34

82

498

r

P

2/28/29

27.2

36.3

8.21

34

83

501

c

n

3/ 2/29

27.5

36.3

8.24

29

83

502

SO

oc

n

3/ 2/29

27.4

36.5

8.24

25

83

503

r

P

3/ 2/29

27.5

36.3

8.24

29

84

SOS

oc

n

3/ 4/29

27.8

36.2

8.23

24

84

506

SO

oc

n

3/ 4/29

27.5

36.4

8.21

24

85

510

oc

n

3/ 6/29

27.9

36.2

8.22

40

85

511

50

r

n

3/ 6/29

27.8

36.2

8.22

40

86

515

.oc

n

3/ 8/29

(28.3

36.2

8.29

20)

86

516

oc

n

3/ 9/29

28.3

36.2

8.29

20

87

521

r

n

3/11/29

27.8

36.1

8.28

17

88

526

oc

n

3/21/29

28.5

35.9

8.23

16

88

527

SO

r

n

3/21/29

28.4

35.9

8.25

13

89

528

oc

n

3/23/29

28.4

35.6

8.25

21

89

529

SO

r

n

3/23/29

28.6

35.8

8.27

12

89

530

r

p

3/23/29

28.4

35.6

8.25

21

89

532

r

n

3/23/29

(28.4

35.6

8.25

21)

90

533

c

n

3/25/29

28.5

35.5

8.27

21

90

534

so

oc

n

3/25/29

28.6

35.6

8.26

21

90

535

oc

P

3/25/29

28.5

35.5

8.27

21

91

540

c

n

3/27/29

28.7

35.1

8.30

21

91

541

50

c

n

3/27/29

28.5

35.2

8.30

24

91

542

r

P

3/27/29

28.7

35.1

8.30

21

92

545

oc

n

3/29/29

28.5

35.3

8.29

28

92

546

SO

c

n

3/29/29

28.4

35.4

8.29

28

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

550

oc

n

3/31/29

28.7

34.7

8.30

28

93

551

50

c

n

3/31/29

28.5

34.8

8.30

28

93

552

r

P

3/31/29

28.7

34.7

8.30

28

94

558

50

r

n

4/22/29

29.3

34.7

8.25

14

94

559

100

r

n

4/22/29

28.5

35.6

8.21

25

95

563

50

c

n

4/24/29

29.3

34.9

8.24

16

95

564

100

oc

n

4/24/29

28.5

35.4

8.22

21

95

566

50

r

P

4/24/29

29.3

34.9

8.24

16

95

568A

oc

n

4/25/29

(29.4

34.7

8.24

14)

96

568

c

n

4/26/29

29.3

35.3

8.26

12

96

569

50

oc

n

4/26/29

29.2

35.3

8.23

12

96

570

100

oc

n

4/26/29

28.2

35.7

8.19

25

96

571

oc

p

4/26/29

29.3

35.3

8.23

12

96

572

SO

r

P

4/26/29

29.2

35.3

8.23

12

96

573

100

oc

P

4/26/29

28.2

35.7

8.19

25

97

574

c

n

4/28/29

28.3

35.2

8.16

24

97

575

50

c

n

4/28/29

28.0

35.4

8.16

21

97

576

100

c

n

4/28/29

27.6

35.6

8.15

25

97

578

50

oc

p

4/28/29

28.0

35.4

8.16

21

97

579

100

r

P

4/28/29

27.6

35.6

8.15

25

98

581

oc

n

4/30/29

27.0

35.3

8.16

24

98

582

50

oc

n

4/30/29

26.9

35.3

8.16

28

98

583

100

oc

n

4/30/29

26.7

35.4

8.14

32

98

585

50

r

P

4/30/29

26.9

35.3

8.16

28

98

588

100

r

P

4/30/29

26.7

35.4

8.14

32

99

589

c

n

5/ 2/29

27.9

34.9

8.21

12

99

590

50

c

n

5/ 2/29

27.8

34.9

8.22

12

99

591

100

c

n

5/ 2/29

27.8

35.0

8.22

17

99

592

oc

P

5/ 2/29

27.9

34.9

8.21

12

99

593

so

oc

P

5/ 2/29

27.8

34.9

8.22

12

99

595

c

n

5/ 2/29

27.9

34.9

8.21

12

100

596

so

c

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

c

n

5/ 4/29

27.6

34.7

8.22

12

100

598

oc

P

5/ 4/29

27.7

34.7

8.21

10

100

599

50

oc

P

5/ 4/29

27.6

34.7

8.21

10

100

600

100

r

P

5/ 4/29

27.6

34.7

8.22

12

100

602

100

r

p

5/ 4/29

27.6

34.7

8.22

12

101

604

50

oc

n

5/ 7/29

26.2

34.7

8.24

8

101

60S

100

r

n

5/ 7/29

25.2

35.1

8.23

8

102

609

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

610

50

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

611

100

oc

n

5/ 9/29

25.6

35.0

8.23

8

103

615

oc

n

5/11/29

26.0

35.0

8.25

5

88

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 25. Distributional and environmental records for Ceratium breve — Concluded

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4

mg/m^

103

617

100

104

621

105

627

50

105

628

50

105

629

100

105

630

106

633

106

635

100

106

636

107

639

107

640

50

107

641

100

107

642

108

646

108

648

100

Guam

652

109

654

109

655

50

111

668

112

674

112

676

100

113

680

113

681

50

113

682

100

113

683

137

836

139

849

139

851

100

140

857

50

140

858

100

140

860

140

861

100

141

864

141

865

50

141

868

141

869

50

149

920

149

921

50

150

929

150

930

50

150

931

100

151

937

151

939

100

151

941

151

942

50

152

944

152

945

50

152

948

153

951

153

952

50

153

953

100

153

955

153

956

50

154

958

154

959

50

154

960

100

155

965

155

966

50

155

967

100

156

972

156

974

100

157

978

157

979

50

157

980

100

158

983

158

984

50

158

985

100

159

990

159

991

50

159

992

100

160

1000

160

1002

50

r

r

c

oc

oc

r

oc

c

r

oc

oc

oc

r

oc

r

oc

oc

oc

r

c

r

oc

oc

oc

oc

r

r

r

r

oc

r

r

r

r

r

r

r

r

c

oc

r

r

r

r

r

oc

oc

r

oc

c

oc

r

r

c

oc

oc

oc

r

oc

oc

oc

c

oc

r

c

oc

oc

oc

r

r

r

r

n

5/11/29

24.8

n

5/13/29

26.1

n

5/15/29

26.8

n

5/15/29

26.8

n

5/15/29

25.2

P

5/15/29

26.9

n

5/17/29

27.2

n

5/17/29

25.6

P

5/17/29

27.2

n

5/19/29

28.0

n

5/19/29

27.9

n

5/19/29

26.8

P

5/19/29

28.0

n

5/27/29

28.4

n

5/27/29

25.2

n

5/ /29

n

5/29/29

27; 4

n

5/29/29

23.1

n

6/ 3/29

20.1

n

6/ 5/29

23.2

n

6/ 5/29

19.8

n

6/25/29

24.2

n

6/25/29

23.8

n

6/25/29

21.5

P

6/25/29

24.2

n

9/18/29

25.5

n

9/22/29

26.7

n

9/22/29

22.4

n

10/ 3/29

26.9

n

10/ 3/29

25.5

P

10/ 3/29

26.9

P

10/ 3/29

25.5

n

10/ 5/29

25.9

n

10/ 5/29

25.9

P

10/ 5/29

25.9

P

10/ 5/29

24.8

n

10/21/29

23.5

n

10/21/29

23.3

n

10/23/29

25.6

n

10/23/29

22.8

n

10/23/29

19.6

n

10/26/29

26.0

n

10/26/29

12.5

P

10/26/29

26.0

P

10/26/29

18.3

n

10/26/29

12.5

n

10/27/29

14.2

P

10/27/29

27.4

n

10/29/29

28.1

n

10/29/29

28.1

n

10/29/29

20.5

P

10/29/29

28.1

p

10/29/29

28.1

n

10/31/29

28.3

n

10/31/29

28.2

n

10/31/29

25.3

n

11/ 2/29

27.8

n

11/ 2/29

27.7

n

11/ 2/29

27.2

n

11/ 4/29

27.6

n

11/ 4/29

26.4

n

11/ 6/29

27.1

n

11/ 6/29

27.1

n

11/ 6/29

26.8

n

11/ 8/29

28.2

n

11/ 8/29

28.2

n

11/ 8/2?

27.6

n

11/11/29

28.6

n

11/11/29

28.5

n

11/11/29

28.0

n

11/13/29

28.6

n

11/13/29

28.6

35.2

8.25

5

35.2

8.24

7

34.9

8.23

5

34.9

8.23

5

35.1

8.23

5

34.9

8.23

5

35.0

8.23

5

35.1

8.23

5

35.0

8.23

5

34.4

8.23

5

34.4

8.23

4

34.9

8.23

11

34.4

8.23

5

35.0

8.25

4

35.0

8.23

4

35!b

8.23

"3

35.0

8.22

3

34.5

8.18

5

34.6

8.22

7

34.7

8.20

8

34.5

8.25

5

34.6

8.25

5

34.7

8.23

8

34.5

8.25

5

35.0

8.39

4

34.8

8.34

6

35.2

8.28

6

35.0

8.39

7

35.0

8.34

7

35.0

8.42

7

35.0

8.34

7

35.2

8.34

5

35.2

8.34

5

35.2

8.34

5

35.3

8.34

5

35.0

8.34

6

35.0

8.37

6

34.7

8.39

7

34.8

8.35

10

34.6

8.32

11

34.0

34.6

34.0

34.4

34.6

34.5

7.87

53

33.7

8.35

20

34.2

8.47

7

34.4

8.39

7

34.7

8.28

31

34.2

8.47

7

34.4

8.39

7

34.2

8.39

7

34.2

8.40

7

34.8

7.93

21

34.9

8.29

29

34.9

8.30

30

35.0

8.30

35

35.0

8.34

28

35.1

8.30

48

35.3

8.27

47

35.2

8.32

60

35.5

8.30

64

35.6

8.34

36

35.6

8.39

50

35.9

8.39

48

35.7

8.37

15

35.7

8.39

15

35.7

8.37

23

35.6

8.37

12

35.6

8.39

15

APPENDIX

8«

Table 26. Distributional and environmental records for Ceratlum euarcuatum

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinit}

PH

P04

(m)

abundance

ratus

(°C)

(o/oo)

mg/m3

1

1

r

n

5/12/28

24.0

36.2

8.16

34

lb

7

50

r

n

5/16/28

22.2

36.4

8.21

39

IS

102

100

r

n

8/11/28

18.4

36.4

8.20

19

16

103

r

p

8/13/28

25.9

36.2

8.24

8

16

104

50

r

P

8/13/28

24.4

36.4

8.23

8

16

107

50

r

n

8/13/28

24.4

36.4

8.23

8

18

118

50

oc

n

8/17/28

22.4

36.8

8.24

5

19

121

50

oc

P

8/20/28

25.2

37.1

8.27

5

19

123

50

oc

n

8/20/28

25.2

37.1

8.27

5

19

124

100

r

n

8/20/28

22.4

37.0

8.25

5

20

125

r

P

8/22/28

26.0

36.6

8.37

5

20

126

50

c

P

8/22/28

25.8

36.6

8.26

3

20

128

r

n

8/22/28

26.0

36.6

8.37

5

20

129

50

oc

n

8/22/28

25.8

36.6

8.26

3

20

130

100

oc

n

8/22/28

22.6

36.7

8.19

5

21

136

50

r

n

8/24/28

24.4

36.2

8.26

4

21

137

100

r

n

8/24/28

21.0

36.8

8.20

7

22

143

50

r

n

8/27/28

24.5

36.2

8.21

9

22

144

100

r

n

8/27/28

17.5

36.1

7.99

123

25

161

50

r

n

9/ 3/28

21.5

36.0

8.22

12

25

162

100

r

n

9/ 3/28

14.6

35.7

7.93

121

26

163

r

p

9/ 5/28

27.6

36.0

8.30

5

27

168

50

oc

p

9/ 7/28

26.0

36.2

8.30

4

27

171

50

r

n

9/ 7/28

26.0

36.2

8.30

4

27

172

100

r

n

9/ 7/28

17.7

36.0

8.09

46

28

174

50

r

p

9/11/28

26.7

36.3

8.26

4

28

183

50

oc

n

9/11/28

26.7

36.3

8.26

4

29

184

100

r

n

9/13/28

23.1

36.6

8.21

8

30

186

50

oc

P

9/15/28

27.8

36.1

8.29

3

30

189

50

oc

n

9/15/28

27.8

36.1

8.29

3

30

190

100

oc

n

9/15/28

24.1

36.4

8.10

20

43

246

100

r

n

11/15/28

13.6

35.0

7.90

92

45

256

oc

n

11/19/28

22.4

35.3

8.12

38

46

264

r

P

11/21/28

23.3

35.3

8.16

36

46

265

50

oc

p

11/21/28

23.2

35.3

8.16

40

48

276

100

r

p

11/25/28

22.7

36.3

8.26

16

49

278

50

r

n

11/27/28

22.6

36.1

8.26

13

49

281

50

r

p

11/27/28

22.6

36.1

8.26

13

50

285

50

r

n

11/29/28

22.0

35.9

8.23

13

50

286

100

oc

n

11/29/28

20.5

35.7

8.22

13

50

288

50

oc

p

11/29/28

22.0

35.9

8.23

13

51

295

100

oc

n

12/ 1/28

20.0

35.6

8.22

17

51

297

50

r

P

12/ 1/28

20.5

35.6

8.22

17

54

320

50

oc

n

12/14/28

19.8

35.4

8.18

17

54

321

100

oc

n

12/14/28

18.7

35.4

8.16

20

55

323

r

n

12/16/28

20.4

34.9

8.19

12

55

325

100

oc

n

12/16/28

16.7

34.9

7.17

12

56

329

50

oc

n

12/18/28

18.5

35.1

8.14

9

56

330

100

c

n

12/18/28

16.6

34.8

8.11

12

56

332

50

oc

p

12/18/28

18.5

35.1

8.14

9

56

333

100

r

p

12/18/28

16.6

34.8

8.11

12

56-57

334

oc

n

12/19/28

19.0

34.5

8.14

20

57

336

50

oc

n

12/20/28

15.6

34.3

8.14

21

57

337

100

oc

n

12/20/28

14.3

34.4

8.10

40

57

338

r

p

12/20/28

19.0

34.5

8.14

20

57

339

50

r

p

12/20/28

15.6

34.3

8.14

21

77

472

r

n

2/18/29

23.7

36.0

8.19

16

77

473

r

p

2/18/29

23.7

36.0

8.19

16

77

474

50

c

p

2/18/29

23.5

36.0

8.19

16

78

476

50

r

n

2/20/29

23.8

36.1

8.14

32

78

477

100

oc

n

2/20/29

21.9

36.2

8.14

34

78

479

50

r

p

2/20/29

23.8

36.1

8.14

34

81

492

50

oc

n

2/26/29

26.4

35.9

8.19

38

81

494

50

r

p

2/26/29

26.4

35.9

8.19

38

81

495

100

r

p

2/26/29

23.6

36.2

8.18

36

82

497

SO

oc

n

2/28/29

27.2

36.3

8.21

34

82

500

100

r

p

2/28/29

24.4

36.3

8.19

34

83

502

SO

oc

n

3/ 2/29

27.4

36.5

8.24

25

83

504

50

r

p

3/ 2/29

27.4

36.5

8>24

25

84

506

SO

oc

n

3/ 4/29

27.5

36.4

8.21

24

84

508

50

r

p

3/ 4/29

27.5

36.4

8.21

24

85

511

SO

r

n

3/ 6/29

27.8

36.2

8.22

40

85

513

SO

r

p

3/ 6/29

27.8

36.2

8.22

40

86

517

50

r

n

3/ 9/29

27.4

36.2

8.29

17

86

519

SO

r

p

3/ 9/29

27.4

36.2

8.29

17

87

525

100

r

P

3/11/29

23.9

36.0

8.23

20

90

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 26. Distributional and enTlronmental records for Ceratium euarcuatum — Continued

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

88

527

50

r

n

3/21/29

28.4

35.9

8.25

13

89

529

50

00

n

3/23/29

28.6

35.8

8.27

12

89

531

100

r

p

3/23/29

26.4

36.0

8.24

12

91

541

50

oc

n

3/27/29

28.5

35.2

8.30

24

92

546

50

oc

n

3/29/29

28.4

35.4

8.29

28

92

548

50

r

p

3/29/29

28.4

35.4

8.29

28

93

551

50

oc

n

3/31/29

28.5

34.8

8.30

28

93

553

100

r

p

3/31/29

27.6

35.8

8.27

29

95

563

50

oc

n

4/24/29

29.3

34.9

8.24

16

95

564

100

oc

n

4/24/29

28.5

35.4

8.22

21

95

566

50

r

p

4/24/29

29.3

34.9

8.24

16

95

567

100

r

p

4/24/29

28.5

35.4

8.22

21

96

569

50

oc

n

4/26/29

29.2

35.3

8.23

12

98

583

100

oc

n

4/30/29

26.7

35.4

8.14

32

98

588

0-100

oc

p

4/30/29

26.7

35.4

8.14

32

99

589

oc

n

5/ 2/29

27.9

34.9

8.21

12

99

592

r

p

5/ 2/29

27.9

34.9

8.21

12

99

595

r

n

5/ 2/29

27.9

34.9

8.21

12

100

596

50

oc

n

5/ 4/29

27.6

34.7

8.21

10

100

597

100

oc

n

5/ 4/29

27.6

34.7

8.22

12

101

604

50

r

n

5/ 7/29

26.2

34.7

8.24

8

102

609

oc

n

5/ 9/29

25.8

35.0

8.24

8

102

610

50

oc

n

5/ 9/29

25.8

35.0

8.24

8

103

615

r

n

5/11/29

26.0

35.0

8.25

5

103

616

50

r

n

5/11/29

25.8

35.2

8.25

5

103

617

100

oc

n

5/11/29

24.8

35.2

8.25

5

104

622

50

oc

n

5/13/29

25.8

35.2

8.24

7

104

623

100

oc

n

5/13/29

25.3

35.3

8.21

7

104

626

100

r

p

5/13/29

25.3

35.3

8.21

7

105

628

50

r

n

5/15/59

26.8

34.9

8.23

5

105

629

100

oc

n

5/15/29

25.2

35.]

8.23

5

106

633

r

n

5/17/29

27.2

35.0

8.23

5

106

634

50

oc

n

5/17/29

27.0

35.0

8.23

5

106

635

100

c

n

5/17/29

25.6

35.1

8.23

5

106

637

50

r

P

5/17/29

27.0

35.0

8.23

5

106

638

100

r

p

5/17/29

25.6

35.1

8.23

S

107

641

100

oc

n

5/19/29

26.8

34.9

8.23

n

108

647

50

oc

n

5/27/29

26.8

35.0

8.24

4

108

648

100

c

n

5/27/29

25.2

35.0

8.23

4

108

650

50

r

P

5/27/29

26.8

35.0

8.24

4

109

656

100

r

n

5/29/29

19.4

34.8

8.18

5

109

658

50

oc

P

5/29/29

23.1

35.0

8.22

3

110

662

50

r

n

5/31/29

18.4

34.8

8.16

7

112

675

50

r

n

6/ 5/29

21.7

34.6

8.23

7

113

680

oc

n

6/25/29

24.2

34.5

8.25

5

132

798

50

oc

n

9/ 8/29

17.6

33.9

8.33

19

132

799

100

r

n

9/ 8/29

14.3

33.4

8.30

16

133

804

oc

n

9/10/29

22.7

34.7

8.47

7

133

805

50

oc

n

9/10/29

20.8

34.7

8.37

7

133

806

100

oc

n

9/10/29

18.4

34.8

8.31

7

133

813

50

r

p

9/10/29

20.8

34.7

8.37

7

133

814

100

r

p

9/10/29

18.4

34.8

8.31

7

133

812

r

p

9/10/29

22.7

34.7

8.47

7

134

808

oc

n

9/12/29

22.9

34.7

8.34

6

134

809

50

oc

n

9/12/29

19.8

34.6

8.34

6

135

821

50

oc

n

9/14/29

21.5

35.0

8.37

5

135

822

100

r

n

9/14/29

18.7

34.8

8.34

5

135

824

r

p

9/14/29

23.8

35.1

8.37

7

135

825

50

r

p

9/14/29

21.5

35.0

8.37

5

136

828

50

c

n

9/16/29

21.4

35.1

8.39

3

136

829

100

c

n

9/16/29

18.6

35.0

8.39

3

136

833

50

r

p

9/16/29

21.4

35.1

8.39

3

137

836

oc

n

9/18/29

25.6

35.0

8.39

4

137

837

50

oc

n

9/18/29

24.4

35.1

8.34

4

138

844

50

r

n

9/18/29

25.6

34.7

8.30

3

138

848

50

oc

P

9/20/29

25.6

34.7

8.30

3

139

850

50

r

n

9/22/29

25.8

34.9

8.31

6

140

857

50

r

n

10/ 3/29

26.9

35.0

8.39

7

141

864

r

n

10/ 5/29

25.9

35.2

8.34

5

141

865

50

oc

n

10/ 5/29

24.8

35.3

8.34

5

141

869

50

r

P

10/ 5/29

24.8

35.3

8.34

5

142

873

50

c

n

10/ 7/29

21.8

34.8

8.30

5

142

874

100

c

n

10/ 7/29

16.6

34.4

8.27

7

142

877

50

r

p

10/ 7/29

21.8

34.8

8.30

5

143

881

r

n

10/ 9/29

22.4

34.4

8.30

6

143

882

50

oc

n

10/ 9/29

22.4

34.4

8.30

6

APPENDIX

91

Table 26. Distributional and environmental records for Ceratlum euarcuatum- -Concluded

Station

Sample

Depth

Relative

Appa-

Date

Temperature

Salinity

pH

P04

mg/m-'

ha^ ^C^ V A^^ll

(m)

abundance

ratus

(°C)

(o/oo)

f**

143

883

100

c

n

10/ 9/29

13.8

34.1

8.30

10

143

855

50

r

P

10/ 9/29

19.0

34.2

8.34

6

144

886

c

n

10/ 9/29

23.3

35.0

8.37

6

144

887

00

P

10/11/29

23.3

35.0

8.37

6

144

888

50

r

P

10/11/29

21.1

34.7

8.33

6

145

891

SO

c

n

10/13/29

18.7

34.3

8.34

6

145

894

50

oc

P

10/13/29

18.7

34.3

8.34

6

145

895

100

r

P

10/13/29

16.0

34.1

8.31

6

146

897

50

oc

n

10/15/29

22.4

34.9

8.30

6

147

903

r

n

10/17/29

23.3

35.3

8.26

8

147

904

50

oc

n

10/17/29

23.1

35.3

8.29

5

148

912

100

oc

n

10/19/29

20.0

35.0

149

921

50

oc

n

10/21/29

23.3

35.0

8.37

6

150

929

r

n

10/23/29

25.6

34.7

8.39

7

150

930

50

r

n

10/23/29

22.8

34.8

8.35

10

150

931

100

oc

r

10/23/29

19.6

3^.6

8.32

11

153

952

50

r

n

10/29/29

28.1

34.4

8.39

7

153

956

50

r

p

10/29/29

28.1

34.4

8.39

7

154

958

r

n

10/31/29

28.3

34.2

8.39

7

154

959

50

r

n

10/31/29

28.2

34.2

8.40

7

155

965

oc

n

11/ 2/29

27.8

34.9

8.29

29

156

972

r

n

11/ 4/29

27.6

35.0

8.34

28

156

967

100

oc

n

11/ 4/29

26.4

35.1

8.30

48

156

973

50

r

n

11/ 4/29

27.0

35.1

8.37

46

156

974

100

r

n

11/ 4/29

26.4

35.1

8.30

48

157

978

oc

n

11/ 6/29

27.1

35.3

8.27

47

157

979

50

oc

n

11/ 6/29

27.1

35.2

8.32

60

157

980

100

oc

n

11/ 6/29

26.8

35.3

8.30

64

158

983

c

n

11/ 8/29

28.2

35.6

8.34

36

158

985

100

r

n

11/ 8/29

27.6

35.9

8.39

48

159

991

50

r

n

11/11/29

28.5

35.7

8.39

15

159

992

100

r

n

11/11/29

28.0

35.7

8.37

23

160

1002

50

r

n

11/13/29

28.6

35.6

8.39

15

Table 27. Distributional and environmental records for Ceratium filicorne

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

P04
mg/m 3

48

276

100

oc

P

49

278

50

oc

n

50

286

100

c

n

50

289

100

oc

p

50-51

290

r

n

51

295

100

c

n

51

298

100

oc

p

52

301

100

c

n

53

305

100

c

n

54

321

100

oc

n

55

325

100

oc

n

56

330

100

c

n

56

333

100

oc

p

57

337

100

c

n

57

340

100

oc

p

63

380

50

c

n

63

381

100

r

n

63

383

50

oc

p

63

384

100

r

p

64

396

100

oc

n

64

399

50

r

p

64

400

100

oc

p

65

406

50

oc

n

66

412

100

oc

n

87

525

100

r

p

91

544

100

oc

p

93

553

100

r

P

11/25/28

11/27/28

11/29/28

11/29/28

11/30/28

12/ 1/28

12/ 1/28

12/ 3/28

12/ 5/28

12/14/28

12/16/28

12/18/28

12/18/28

12/20/28

12/20/28

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 3/29

1/ 3/29

1/ 3/29

1/ 5/29

1/ 7/29

3/11/29

3/27/29

3/31/29

22.7

36.3

8.26

16

22.6

36.1

8.26

13

20.5

35.7

8.22

13

20.5

35.7

8.22

13

(22.8

35.6

8.22

16)

20.0

35.6

8.22

16

20.0

35.6

8.22

16

18.2

35.2

8.17

8

19.9

35.6

8.19

13

18.7

35.4

8.16

20

16.7

34.9

7.17

12

16.6

34.8

8.11

12

16.6

34.8

8.11

12

14.3

34.4

8.10

40

14.3

34.4

8.10

40

17.0

34.6

8.08

25

15.6

34.6

8.08

25

17.0

34.6

8.08

25

15.6

34.6

8.08

24

15.8

34.5

8.10

32

17.2

34.6

8.12

29

15.8

34.5

8.10

32

16.5

34.5

8.10

25

17.8

34.9

8.12

21

23.9

36.0

8.23

20

25.8

36.0

8.25

30

27.6

35.8

8.27

29

92

CERATIUM IN THE PACIFIC AND NORTH ATLANTIC OCEANS

Table 28. Distributional and environmental records for Ceratium symmetricum

Station

Sample

Depth
(m)

Relative
abundance

Appa-
ratus

Date

Temperature
(°C)

Salinity
(o/oo)

pH

PO4
mg/m3

Variety

la

4

la

5

lb

7

50

lb

8

100

15

99

100

IS

102

100

16

104

50

16

107

50

16

108

100

17

112

50

17

113

100

18

116

100

18

118

50

18

119

100

19

123

50

19

124

100

20

130

100

23

146

50

23

149

SO

24

152

50

25

161

50

26

165

100

27

172

100

29

179

29

180

50

29

183

50

32

196

100

35-36

211

36

214

50

37

218

50

41

239

100

42

243

100

45

257

50

45

258

100

45

259

45

260

SO

46

261

46

262

50

46

263

100

46

265

SO

47

266

47

267

SO

47

268

100

47

270

50

48

274

48

275

50

48

277

49

278

SO

51

294

50

51

295

100

56

330

100

57

337

100

57

340

100

58

346

100

59

352

50

60

354

62

373

100

62

376

100

63

380

50

63

382

63

383

50

63

384

100

63-64

387

63-64

388

63-64

389

63-64

390

63-64

392

64

395

50

64

398

64

399

50

64

400

100

64-65

401

65

406

50

65

407

100

66

412

100

66

415

100

r

r

oc

r

r

p

r

r

r

r

oc

r

r

c

oc

oc

r

r

r

r

r

r

r

r

oc

oc

r

r

r

r

r

r

c

oc

oc

p

oc

r

r

oc

p

oc

oc

p

p

p

p

p

oc

oc

c

oc

r

r

r

r

p

r

c

oc

oc

p

oc

p

r

r

oc

p

p

p

oc

r

oc

p

oc

r

n
n
n
n

P
n

P
n
n
n
n

P
n
n
n
n
n

P
n

P
n

P
n

P
P
n
n
n
n
n

P
n
n
n
P
P
n
n
n

P
n
n
n
P
P
P
n
n
n
n
n
n
P
P
P
n
n

P
n

P
P
P
n
n
n
n
n
n
P
P
P
n
n
n
n
P

5/14/28

5/14/28

5/16/28

5/16/28

8/11/28

8/11/28

8/13/28

8/13/28

8/13/28

8/15/28

8/15/28

8/17/28

8/17/28

8/17/28

8/20/28

8/20/28

8/22/28

8/29/28

8/29/28

8/31/28

9/ 3/28

9/ 5/28

9/ 7/28

9/13/28

9/13/28

9/13/28

10/ 5/28

10/28/28

10/30/28

11/ 1/28

11/10/28

11/13/28

11/13/28

11/19/28

11/19/28

11/19/28

11/21/28

11/21/28

11/21/28

11/21/28

11/23/28

11/23/28

11/23/28

11/23/28

11/25/28

11/25/28

11/25/28

11/27/28

12/ 1/28

12/ 1/28

12/20/28

12/20/28

12/20/28

12/22/28

12/24/28

12/26/28

12/30/28

12/30/28

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 1/29

1/ 3/29

1/ 3/29

1/ 3/29

1/ 3/29

1/ 3/29

1/ 5/29

1/ 5/29

1/ 7/29

1/ 7/29

122.3

36.3

8.20

46)

or

22.3

36.3

8.20

46)

or

22.2

36.4

8.21

39)

or

20.3

35.5

8.18

36)

sy

18.4

36.4

8.20

19

or

18.4

36.4

8.20

19

or

24.4

36.4

8.23

8

CO

24.4

36.4

8.23

8

CO

19.9

36.5

8.17

13

CO

21.9

36.6

8.28

12

or

19.3

36.5

8.23

9

or

20.4

36.8

8.21

5

CO

22.4

36.8

8.24

5

CO

20.4

36.8

8.21

5

CO

25.2

37.1

8.27

5

CO

22.4

37.0

8.25

5

CO

22.6

36.7

8.19

5

sy

20.9

36.0

8.14

13

CO

20.9

36.0

8.14

13

CO

23.1

36.0

8.14

8

CO

21.5

36.0

8.22

12

CO, sy

14.9

35.6

8.11

40

CO

17.7

36.0

8.09

46

CO

27.6

36.2

8.31

8

or

27.2

36.2

8.29

3

or

27.2

36.2

8.29

3

or

22.2

36.4

8.10

30

CO

(27.0

30.7

8.27

16)

sy

18.5

34.5

8.03

122

sy

18.8

34.5

8.00

121

CO

14.5

35.0

7.91

152

CO

13.8

35.0

7.91

150

CO

17.2

34.9

7.99

68

or

18.6

35.1

8.00

50

or

22.4

35.3

8.12

38

or

22.4

35.2

8.13

46

or

23.3

35.3

8.16

36

or

23.2

35.3

8.16

40

or

22.5

35.4

8.17

40

CO

23.2

35.3

8.16

40

or

23.9

36.0

8.23

17

or

23.8

36.0

8.23

20

CO, or

22.7

36.2

8.23

20

CO, or

23.8

36.0

8.23

20

or

23.6

36.4

8.23

13

or

23.6

36.4

8.24

16

or

23.6

36.4

8.23

13

or

22.6

36.1

8.26

13

or

20.5

35.6

8.22

17

or

20.0

35.6

8.22

17

or

16.6

34.8

8.11

12

CO

14.3

34.4

8.10

40

or

14.3

34.4

8.10

40

sy

12.3

34.1

8.05

40

sy

14.0

34.0

8.08

38

sy

15.0