DEPARTMENT OF TERRESTRIAL MAGNETISM

J. A. Fleming, Director

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

BIOLOGY- II

The Oceanic Tintinnoina of the Plankton
Gathered during the Last Cruise

of the Carnegie

ARTHUR SHACKLETON CAMPBELL

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I LIBRARY

CARNEGIE INSTITUTION OF WASHINGTON PUBLICATION 537

WASHINGTON, D. C.
1942

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DEPARTMENT OF TERRESTRIAL MAGNETISM

J. A. Fleming, Director

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

BIOLOGY- 1 1

The Oceanic Tintinnoina of the Plankton
Gathered during the Last Cruise

of the Carnegie

ARTHUR SHACKLETON CAMPBELL

* — 1R23»vi

LIBRARY

%$V».

CARNEGIE INSTITUTION OF WASHINGTON PUBLICATION 537

WASHINGTON, D. C.
1942

This book first issued September 18, 1942

^ <z - <J

THE WILLIAM BYRD PRESS, RICHMOND, VIRGINIA

THE MERIDEN CRAVURE COMPANY, MERIDEN, CONNECTICUT

THE VIRGINIA ENGRAVING COMPANY, RICHMOND, VIRGINIA

PREFACE

Of the tio,000 nautical miles planned for the seventh
cruise of the nonmagnetic ship Carnegie of the Carnegie
Institution of Washington, nearly one-hall had been com-
pleted upon her arrival at Apia, November 28, [929. The
extensive program of observation in terrestrial mag-
netism, 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. In addition to the extensive magnetic and
atmospheric-electric observations, a great number of data
and marine collections had been obtained in the fields of
chemistry, physics, and biology, including bottom samples
and depth determinations. These observations were made
at 162 stations, at an average distance apart of 300 nautical
miles. The distribution of these stations is shown in map
1, which delineates also the course followed by the vessel
from Washington, May 1, 192S, to Apia, November 28,
1929. At each station, salinities and temperatures were
obtained at depths ol 0, 5, 25, 50, 75, too, 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 samples
to the number of 1014 were obtained both by net and by
pump, usually at 0, 50, and 100 meters. Numerous phys-
ical 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 accordance
with the established policy of the Department of Ter-
restrial Magnetism, all observational data and materials
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 re-
sults obtained during this last cruise of the Carnegie are
being published under the classifications Physical Ocean-
ography, Chemical Oceanography, Meteorology, and
Biologv, in a series numbered, under each subject, 1, II,
111. etc.

A general account of the expedition has been prepared
and published by J. Harland Paul, ship's surgeon and
observer, 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 equipment,
and a full narrative of the cruise (Baltimore, Williams
and Wilkins Company, 1932; xiii -f 331 pages with to8
illustrations).

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 Depart-
ment; 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 Com-
parative Zoology of Harvard University; the School of
Geography of Clark University; the American Radio
Relay League; the Geophysical Institute, Bergen, Nor-
way; the Marine Biological Association of the United
Kingdom, Plymouth, England; the German Atlantic
Expedition of the Meteor, Institut fur Meereskunde, Ber-
lin, Germany; the British Admiralty, London, England;
the Carlsberg Laboratorium, Bureau International pour
I'Exploration de la Mer, and Laboratoire Hydrogra-
phique, 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 Research Associate of
the Carnegie Institution of Washington at the Geophys-
ical Institute at Bergen, Norway, was consulting ocean-
ographer 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 quarter
of a century, and which required cooperative effort and
unselfish interest on the part of many skilled scientists,
it is impossible to allocate full and appropriate credit.

111

IV

PREFACE

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 what Captain Peters had
so well begun. The original plan of the work was en-
visioned by L. A. Bauer, the first Director of the Depart-
ment of Terrestrial Magnetism, Carnegie Institution 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."

The following memoir by Dr. Arthur S. Campbell is
the second in the series of reports on studies of the
plankton gathered during the last cruise of the Carnegie.
Of value in this study of the oceanic Tintinnoina has
been the mass of material accumulated by other vessels,
especially the Albatross and the National, detailed ac-
counts of which are given elsewhere.

J. A. Fleming
Director, Department of Terrestrial Magnetism

CONTENTS

PAGE

Introduction i

Acknowledgments 2

Methods 3

Taxonomy and Distribution of the Tintinnoina

Found in the Material of the Carnegie 6

Codonellidae 6

Cyttarocylidae 17

Codonellopsidae 22

Dictyocystidae 31

Coxliellidae 38

Favellidae 43

Ptychocylidae 46

Petalotrichidae 48

Rhabdonellidae

Epiplocylidae

Xystonellidae

Undellidae . . .
Tintinnidae . .

PARE

• 54

. 65

■ 75

• 94
. in

Summary of Results
Literature Cited

Plate i

Figures 1-128

Systematic Index . . . .

134

'34

jiiiing 136

137

159

THE OCEANIC TINTINNOINA OF THE PLANKTON GATHERED DURING THE

LAST CRUISE OF THE CARNEGIE

INTRODUCTION

The Tintinnoina arc one of the major subdivisions ol
the ciliatc protozoans. The group is among the more
complex of these ciliates, ranking in structural elaboration
with the symbiotic ciliates of ruminant mammals and
the peritrichous ciliates.

The body is conical or trumpet-shaped, and attached
by the inverted apex to an external lorica, which is not in
close contact with the animal as in the Sarcodina anil
Mastigophora. The broad basal end of the body projects
out of the lorica through the oral opening of the latter
and is provided with a crown of highly differentiated
motor organs. These are relatively few in number (12 to
24) and of complicated structure. This wreath or circlet
of membranelles is composed of separate blades, the
whole forming a left-twisted spiral, one end of which
enters a preoral cavity on one side of the depressed
frontal area, in the middle of which arises a piston
organelle. Each blade is made up of a line of 8 adjacent
lamellae, each consisting of a dense rodlet and a thin,
wide, hyaline sheet. In cross section the lamella is sickle-
shaped and presents the following structures: (a) an en-
closing pellicular envelope, (b) a ground substance in
which are embedded 6 fibrils in the blunted outer and
denser part, and a single one in the hyaline material on
the convex side of the lamella. In the blunt end are vari-
cosed alveoles strung along an irregular line. Basal
granules connect this system of fibrils, and these in turn
are in contact with the neuromotorium. On the inner
face of each blade is a lappet organ.

The neuromotorium is a granular spindle-shaped or
even round body lying between the body wall of the ani-
mal and the gullet, which latter is below the preoral de-
pression, and into which extends a brush of cilia. The
neuromotorium is the dynamic center of the animal. It
serves as the center of coordination in locomotion and
other body activity, and at the time of binary fission be-
comes the directing agent for the forming of the lorica.
From the neuromotorium arise several fibers, the most
important of which connects with the membranelles.
Others probably connect the body cilia and the ciliary
membrane when this last structure is present.

The ciliary membrane is present in Tintinnidium, Tin-
tinnopsis, Codonella, Stenosemella, Codonellopsis, and
probably other genera. It is made up of a line of strong,
long cilia which extend down the body on the ventral
(gullet) side. Body cilia are mainly confined to the

upper three-tenths ol the body. They arc sparse and often
rather long. Some arc used as stays by the animal when
extended from the lorica. Small patches of cilia an- often
found in rows below the peristome, on the body.

The animal has 1 to 100 macronuclei, the number
being usually a pair, and 8 to 20 being common in
eupelagic species; the number is constant save during
reproductive periods. The macronuclei are large and
sausage-shaped. The micronuclei are small and globular.
They lie near the macronuclei and are usually similar in
number to the latter.

During reproduction by binary fission, a chromophile
cloud of granules is accumulated in the body in the
region of the gullet. This material is prominent at about
the time the cytoplasmic organs appear. As progressive
nuclear and cytoplasmic changes occur, these granules
come to form a shieldlike mass above the frontal area of
the forming posterior schizont. That this material forms
the substance of the lorica is evident from the following
considerations: (a) its structure is alveolar, (b) its con-
sistency resembles that of the lorica, (c) it makes its
appearance at the time of fission and at no other time,
(d) it occurs in all species examined, (e) it forms in a
position such that it may be discharged by existing
organs, probably the mouth, and the discharged material
is whipped about by the membranelles. These data fit in
well with observations on tintinnids which have been
watched in the living state at the time of fission. In
short, the lorica is the product of secretion and behavior.

The lorica is usually cuplike, with a single large open-
ing. The morphological characters of diagnostic value
are: the finer structure of the wall, which may be laminar,
prismatic, alveolar, or hyaline; the inclusions and the
tendency to incorporate adherent detritus; the propor-
tions, which are generally pretty constant; the suboral
and circumoral differentiations, including the nature of
the margin, the collar, teeth, bands, rings, and ledges, and
the interpolated collars or other structures; the general
proportions of the bowl, its outline, and the extent and
level of its maximum diameter; the aboral structures,
such as the pedicel, knob, lance, apophyses, skirt, and
other significant differentiations, and the form and pro-
portions of the aboral opening when present; the differ-
entiations of the surface of the lorica, such as rings,
angles, ribs, ridges, plications, shelves, and (lutings, to-
gether with the number, spacing, distribution, direction,

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

and anastomoses of these elements, also reticulations,
fenestrae, lacunae, and coccoliths. The actual dimensions
of the lorica are subject to correlation with physical factors
and do not lend themselves to more than general de-
scription.

The lorica is used as the basis of classification, as is
the shell in the Foraminifera, Radiolaria, Dinoflagellata,
Mollusca, and Arthropoda, and the skeleton in the verte-
brates. The outstanding feature in the speciating process
and in generic differentiation in the Tintinnoina is the
play of behavior, of the movements of the differentiating
schizonts, in the distribution of the secreted material and
in its shaping into the generic and specific patterns. This
coordinated behavior on the part of the daughters begins
before the separation and continues beyond the period
of cytoplasmic discontinuity. Thus, the oral and aboral
ends of the lorica, the regions in contact with the proto-
plasm for the longest time at binary fission, are the most
importantly modified and constitute the regions of great-
est taxonomic significance. The specific characters are
largely these, and generic characters are found in the con-
sistency of the wall substance and in the general form.
The oral end is directed backward in locomotion, and
the aboral end meets the first shock of environment. The
former has usually outrun the latter in differentiation,
and this has developed, perhaps, in response to specific
modification in behavior rather than from surrounding
influences.

The Tintinnoina constitute about 40 per cent of the
known ciliates, marine and fresh-water. Few marine
ciliates are nonloricate. Their freedom of movement and
the availability of food in the sea has doubtless led to a
wide evolutionary spread of these creatures. No sea is
without some representative of this suborder. In the
coldest waters of the Arctic and within the ice-bound
Antarctic these animals are fully at home. They reach
their greatest differentiation in kind, however, in the
warm waters of the equatorial regions of the ocean. Here
there are hundreds of kinds in every plankton catch.
Especially within the upper levels of these warm seas,
where Coccolithophoridae are frequent and light pene-
trates freely, do these creatures abound.

Acknowledgments

Acknowledgment is made to the Carnegie Corporation
of New York for funds with which to complete the plate
and the figures. Mr. H. W. Graham, and Dr. J. A.
Fleming, Director of the Department of Terrestrial Mag-
netism of the Carnegie Institution of Washington, gave
help from time to time. Mrs. A. Abernathy assisted in
the execution of the finished drawings. The writer's
wife was an invaluable assistant in many ways, and the
administration of St. Mary's College offered sympathetic
aid. Without all these, the preparation of this report
would have been hindered.

METHODS

The samples were examined in the original sea water
and formalin without other treatment than the occasional
replacement of loss by evaporation with distilled water.
It is thus possible carefully to orient the organism under
the cover slip and to secure diverse views of the same
individual. There does not seem to he any satisfactory
method of sealing off mounts in formalin. A few smears
of plankton were made, however, and mounted either in
Venice turpentine or in balsam. These were used to
supplement the routine examinations. Adequate micro-
scopes and maximum equipment were used at all times
in the examination of the material.

After a sufficient knowledge of the species of Tintin-
noina had been obtained, each sample was systematically
searched by the use of the mechanical stage until all the
species had been detected. Each sample, after being
thoroughly shaken, was examined on a standard slide
under a long cover slip, and several slides were made of
it. When the whole sample had been investigated, the
material was washed back into the original phial. Care
was taken that slides, slips, pipettes, and other instru-
ments were cleaned before another sample was opened.
The camera lucida was always attached to the microscope
so that drawings could be made at once when a desired
individual was discovered. In order to have some sem-
blance of quantitative record of frequency in individuals
of the component species of each net sample, records
were made during the search of the sample up to the
first hundred individuals examined. Thereafter each
additional species was recorded as merely present. The
number of individuals recorded is thus the percentage in
that sample. These records refer solely to the relative
numbers of different species in the one sample, and do
not indicate the relative numbers of the same species in
other samples. In the examination of the pump samples,
the whole number of individuals of all species found was
counted out directly, so that in some instances several
hundred were recorded. These are reported fully. In
most cases the numbers were small and are recorded with
those of the net samples without differentiation.

The manner of collection, and of accumulation of
physical data, is recorded in the general survey of the
expedition, as is also the list of species by station and
depth.

Drawings with the camera lucida at standard magni-
fications were accumulated in great number during the
examination of the plankton. Several thousand sketches,
with additional notes when necessary, were thus brought
together. From these the finished drawings were made
up, and the range of variation ascertained. All the

descriptions of species in this report are the result of the
examination and measurement oi these drawings. In ibis
way it became possible to describe a large fauna accu-
rately and to determine at a later time the exact status oi
aberrant individuals. Stress was laid on the finding and
recording of these unusual individuals. Most of the fin-
ished drawings are selected to show some unusual feature
of morphology. Since the majority ol the oceanic spei i< S
have been figured previously, it was not considered neces-
sary to illustrate all species, especially in those instances
in which the material of the Carnegie was not particu-
larly different from that already described. The descrip-
tions in all instances, however, are of loricae of species
found in the collection. Thus these descriptions may
differ from those given in earlier papers.

In this report each character has approximately the
same place in the description and so far as possible is
discussed in similar phraseology. For the families, sub-
families, and genera, synonymy, relationships, and gen-
eral distribution are given. The descriptions of the
species include synonymy, description of specimens found
in the Carnegie material, variations, comparisons, history,
and occurrence in the material of the expedition. Because
the various genera are distinctly different in form, it is
not always possible to apply the same procedure of
description to them all, or to use an identical terminology.
Adjectives and adverbs used in the descriptions are to be
understood in their usual sense, although they are seldom
quantitatively exact.

The families, subfamilies, and genera are arranged in
a presumed order of natural relationship, based on the
comparative structure of the lorica. The species are
arranged within each genus alphabetically. A number of
changes in nomenclature and systematic arrangement
have been made in an attempt more naturally to separate
the families, genera, and species. The relations of the
species to one another are treated in the paragraph on
comparisons.

In the synonymy, only those forms recorded since the
publication of the Conspectus by Kofoid and Campbell
(1929) are given. In that monograph all previous sig-
nificant literature is reviewed.

The records of distribution are arranged according to
the natural areas of distribution of Gerhard Schott, which
regions are based on the physical conditions of the sea
water. The geographical distribution thus does not
strictly follow the ocean currents, since these are difficult
to limit and are subject to different physical conditions
along their lengths. The accompanying table gives a
summary of the route of the Carnegie. See also map 1.

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Leg

Depart-
ing

From

Arriv-
ing

At

Dis-
tance
(miles)

1928-9

1928-9

1

May 1 1

Newport News

June 9

Plymouth

3.669

2

June 18

Plymouth

June 22

Hamburg

614

3

July 7

1 l.imburg

July 20

Reykjavik

1.329

4

July 27

Reykjavik

Sept. 16

Barbados

5.7I.S

5

Oct. 1

Barbados

Oct. 11

Balboa

1,361

6

Oct. 25

Balboa

Dec. 6

Easter Island

4788

7

Dec. 12

Easter Island

Jan. 14

Callao

3,334

8

Feb. 5

Callao

Mar. 13

Papeete

4,470

9

Mar. 20

Papeete

Apr. 1

Pago Pago

1,274

10

Apr. 5

Pago Pago

Apr. 6

Apia

80

1 1

Apr. 20

Apia

May 20

Guam

3,914

12

May 25

Guam

June 7

Yokohama

1,447

13

June 24

Yokohama

July 28

San FYancisco

4,839

14

Sept. 3

San Francisco

Sept. 23-

Honolulu

2,l86

15

Oct. 2

Honolulu

Nov. 18

Pago Pago

5,777

16

Nov. 27

Pago Pago

Nov. 28

Apia

80

Total 44.877

It will be seen that a wider area of the oceans was covered
than has been investigated before by almost any other
single expedition. The wealth of physical data that
accompanies the plankton catches makes this voyage
especially valuable.

The natural areas of distribution covered are these: the
Gulf Stream, the Atlantic drift, the North Sea, the
American cold-water region, the Sargasso Sea, the
Atlantic equatorial region, the Caribbean Sea, the East

Asiatic marginal sea, the Alaskan secondary region, the
North Pacific middle latitudes, the California region, the
North Pacific trade region, the Pacific equatorial region,
the Galapagos region, the region of South Pacific island
fields, and the South Pacific middle latitudes. In each of
these sixteen areas some Tintinnoina were found. In the
accounts of occurrence, the order of the ship's route is
followed.

A total of 648 phials was examined. Of these, 272 were
in net samples and 376 were in pump samples. There-
were 70 net samples from the Atlantic and 202 from the
Pacific. Of pump samples, there were 58 from the
Atlantic and 318 from the Pacific. No net samples were
available from stations 86 to 94 inclusive, as there was no
residual lot remaining at the time this investigation was
in progress. Scattered net samples were also missing
throughout the collection, but pump samples were ob-
tained at these stations. In 51 of the net samples no
Tintinnoina were recorded. These are Carnegie samples
9087, 9089, 9090, 9091, 9093, 9094, 9095, 91 1 1, 9112, 91 1 8,
91 19, 9120, 9121, 9122, 9123, 9129, 9130, 9131, 9143, 9154,
9172, 9175, 9178, 91S1, 9183, 9186, 9202, 9213, 9219, 9220,
9221, 9231, 9234, 9239, 9243, 9248, 9273, 9274, 9284, 9286,
(,1287, 9291, 9312, 9314, 9315, 9317, 9318, 9321, 9329, 9331,
9347. Most of these are from transitional regions. In 37
pump samples no Tintinnoina were recorded: 2672, 2695,
2715, 2726, 2729, 2761, 2766, 2769, 2771, 2774, 2775, 2778,
2780, 2797, 2816, 2818, 2819, 2820, 2835, 2842, 2843, 2853,
2855, 2859, 2865, 2877, 2903, 2918, 2927, 2928, 2929, 2930,
2964, 2966, 2967, 2973, 3163.

TAXONOMY AND DISTRIBUTION OF THE TINTINNOINA FOUND
IN THE MATERIAL OF THE CARNEGIE

(Temperatures are given in degrees centigrade. Salinities are expressed per mille. Values of density are in units of the third
decimal, and express the excess over unity; thus, 23.26 signifies a density of 1.02326. Values of pH are in the usual units

expressing hydrogen-ion concentration.)

CODONELLIDAE Kent emended

Codonellidae, Kofoid and Campbell, 1929, p. 18.

The family includes four genera: Tintinnopsis, Codonclla,
Codonaria, and Codonopsis. These genera are largely ma-
rine, only a few odd species of Tintinnopsis and Codonclla
being found in fresh and brackish water. Most species of
Tintinnopsis and one or two of Codonella occur in coastal
waters; Codonaria and Codonopsis are exclusively eupelagic,
nearly always in warm seas.

All four genera occur in the Carnegie material.

TINTINNOPSIS Stein emended
Tintinnopsis, Kofoid and Campbell, 1929, pp. 19-20.

Species of Tintinnopsis are usually rare in oceanic plank-
ton, the genus commonly occurring in neritic or even
brackish-water conditions. Under the varied environments
of coastal waters, many local species have come into being.
Only a few are found in the high sea and these are appar-
ently developed from species of the coasts. Tintinnopsis is
scarce in warm and temperate water except northward,
where there are a number of coastal forms along the shore
line. The genus does not enter the Antarctic.

Two species are described, of which one is new.

Tintinnopsis penrhynensis, new species
(Plate 1, figure 3)

The tiny lorica has a globose bowl and a ringlike collar,
and its length is 1.56 oral diameters. The oral margin is
irregularly beset with minute blobs of alveolar matter so that
its edge is ragged. The collar is a cufflike tube with a length
of 0.53 oral diameter, and has a scarcely detectable median
bulge with a diameter little greater than that of the oral
opening. The globular bowl reaches its greatest diameter,
1.53 oral diameters, at the middle. The upper, open end
joins the collar, forming a modest shoulder, and the lower
hemisphere rounds out, there being no aboral differentiation.

The wall is 0.14 oral diameter in thickness near its thickest
level, which is across the level of the junction of collar and
bowl; it gradually thins and becomes extraordinarily thin at
the aboral end. There are coarse, irregular, crudely pris-
matic tertiary structures in the wall. The outer surface
shows little irregularity, and few blobs occur.

Length, total 50(1, collar 12.5(1; diameter, oral 32(1, throat

35H-

This is a locally developed species of Tintinnopsis, and
differs from all others in shape. It is somewhat like T. ber-
mudensis in having a cufflikc collar and round bowl, but in

bermudcnsis the collar is much taller and more funnel-like,
and the bowl is pointed. It bears a little resemblance to T.
baltica, but in baltica the collar widens out suborally, and
again the bowl is aborally pointed. It looks like T. bornandi,
but the collar is cufllike, taller, and more regular. It is some-
what like T . conglobata, but is shorter and stouter, with a
rounded instead of ovate bowl, and with a more clearly de-
veloped shoulder; it is not much like other species of
Tintinnopsis described from the coast of Asia or elsewhere
in the Pacific.

Recorded from the lagoon of Penrhyn Island, a surface
catch on November 10, 1929. Only 2 loricae were observed.

No physical data are available for this exceedingly inter-
esting locality.

Type locality, Penrhyn Island lagoon (between stations
159 and 160), at surface; latitude 9 south, longitude 158°
west.

Tintinnopsis rara Kofoid and Campbell

Tintinnopsis rara Kofoid and Campbell, 1939, p. 41, pi. 1. fig. 7.

The stout lorica, with cylindrical collar and globular bowl,
has a length of 1.57 oral diameters. The oral margin is thin,
spreading, and slightly recurved. The low collar is a cylin-
der with a length of 0.3 oral diameter, with laterally concave
sides and somewhat spreading upper and lower parts. The
bowl is practically hemispherical without any special differ-
entiation at the aboral end; its diameter is 1.4 oral diameters.

The wall is coarsely reticulated with large irregular poly-
gons or tertiary prisms, with thickened edges and clear con-
tents. The wall is rather thin, and the cavity follows exactly
the outer contour.

Length, 56(1.

Kofoid and Campbell (1929) called this species Tintin-
nopsis bornandi; the Pacific form is different from the Medi-
terranean species, however, as these writers later recognized.

Tintinnopsis rara is closest to T. bornandi in form and
size, but the surface is not beset with adherent irregular
particles, the bowl is more symmetrically globular, and the
collar is a trifle lower, all these characters possibly being
associated with the more definitely pelagic as contrasted with
neritic habit.

Recorded from three stations in the Pacific, as follows:
one (65) in the South Pacific middle latitudes, one (96) in
the region of South Pacific island fields, and one (13s) in
the California region.

Pump samples only, 2 from 50 meters and 1 from the sur-
face. Frequency, minimum.

Temperature, i6?96-29°3o (22?69); salinity, 34.47-35.27
(34.91); density, 22.19-25.14 (23.87); pH, 8.10-8.37 ( 8 - 2 3)-

TAXONOMY AND DISTRIBUTION

CODONELLA Haeckel emended
Codonella, Kofoid and Campbell, 1929, pp. 51-52.

No fresh-water Tintinnoina were recorded in the material
of this expedition, although 2 samples were taken in Lake
Vehire at Tahiti. In these samples there were diatoms, small
annulates, rotifers, and abundant statoblasts of the bryozoan
Plumatella sp. Codonella cratera might have been expected;
the lack of fresh-water Tintinnoina in mid-Pacific oceanic
island lakes is interesting. Nearly every large body of fresh
water on the continents has some of these ciliates. Codonella
rarely enters coastal waters, being mainly confined to the
warm oceans. No species are found in the Antarctic, and
only a few in cooler northern seas.

Fifteen species are described, of which one is new.

Codonella acerca JSrgensen

Codonella acerca, Kofoid and Campbell, 1929, p. 52, fig. 11 ^.

The short lorica, with funnel-like collar, rotund bowl, and
pointed aboral end, has a length of nearly 1.56 oral diameters.
The oral margin is thin-edged and entire. The collar is a
segment of an inverted, slightly expanded cone (20 ) with
a length of 0.5 oral diameter, and a diameter at the lower
end of 0.81 oral diameter. Its sides are strictly plane. The
plump bowl reaches its greatest diameter, r.12 oral diam-
eters, at 1.0 oral diameter below the rim. The lower bowl
gradually contracts, reaching 1.0 oral diameter at 1.31 oral
diameters below the rim, and then rather more suddenly
becomes a wide inverted convex cone (110°) with an un-
modified, not prolonged point.

The wall is thin, being hardly 0.02 oral diameter in thick-
ness in the bowl. There are feeble laminae with long, rec-
tangular secondary prisms in but one layer. There is a weak
internal ledge. The exterior surface has unequal, round to
polygonal meshes. The large polygons are best formed in
the upper bowl.

Length, 64 to 7211.

Codonella acerca differs from C. nationalis chiefly in the
pointed rather than rounded aboral region.

Recorded from one station (18) in the Sargasso Sea, in a
net sample taken at 100 meters. Frequency, minimum.

Temperature, 20?32; salinity, 36.81; density, 26.06; pH,
8.21.

Codonella acuta Kofoid and Campbell
Codonella acuta Kofoid and Campbell, 1929, p. 52, fig. 104.

The fairly tall lorica, with funnel-like collar, broadly ovate
bowl, and acutely pointed aboral end, has a length of 1.57
oral diameters. The entire, thin-edged oral rim has a hyaline
band with a width of less than 0.1 oral diameter. The in-
verted, funnel-shaped (20°) collar, the length of which is
0.25 total length, reaches a diameter of 0.83 oral diameter at
the neck; sometimes there is a short sigmoid curve in the
wall just above its junction with the bowl. The rather
plump bowl expands from the throat and reaches 1.08 oral
diameters at 0.53 total length from the rim. Below this level
the bowl contracts with full, convex sides to the simple,
obtusely angular (105°) aboral end.

The wall reaches a thickness of 0.06 oral diameter across

the bowl and is much thinner in the collar. There is some-
times a double layer of large, rectangular secondary prisms.
The outer surface has very thick-walled, rounded secondary
polygons in the upper half and larger ones below. Usually
these polygons enclose circular coccoliths and rhabdoliths.

Length, 80 to 1 05U..

The coccolith-bearing habit is well developed in this
species, although some loricac apparently lack them. The
length is variable, owing, no doubt, to temperature relations.

Codonella acuta has a more deeply constricted throat than
C. galea, as well as a pointed aboral end. There is no pro-
jecting point as in C. cuspidata or tropica. It is not so tall
nor so trim as C. elongata, and its bowl is much wider; the
aboral end is acute rather than narrowly rounded.

Recorded I mm sixteen stations in the Pacific, as follows:
three (45, 46, 71) in the Galapagos region, seven (81, 90,
96, 98, 158, 159, 160) in the region of South Pacific island
fields, four (101, 104, 138, 140) in the North Pacific trade
region, one (131) in the California region, and one ( 145) in
the North Pacific middle latitudes.

There are 11 pump and 10 net samples, of which 4 were
taken at the surface, 8 at 50 meters, and 9 at 100 meters.
Maximum frequency, 28 per cent at station 159 at 50 meters;
other records above minimum (2 to 5 per cent) from stations
81, 131, 140, 159, 160; average in net samples, 6.8 per cent.

Temperature: pump samples i6?58-28?5o (24^86), net
samples i2°i2-2j°^o (23?42). Salinity: pump samples
34.18-35.82 (35.18), net samples 33.30-35.89 (35.31). Den-
sity: pump samples 22.62-25.36 (23.44), net samples 22.08-
25.31 (23.67). pH: pump samples 7.96-8.34 (8.21), net
samples 8.12-8.39 (^- 2 7)-

Codonella amphorella Biedermann
(Figure 11)
Codonella amphorella, Kofoid and Campbell, 1929, p. =;s. fig-
132.

The short, plump lorica, with convex collar, egg-shaped
bowl, and short aboral horn, has a length of 1.8 oral diam-
eters. The oral margin is smooth-edged and thin-lipped.
The swollen collar has a length of 0.36 oral diameter, and a
greatest diameter of 1.12 oral diameters just a little above its
middle. The throat attains 0.96 oral diameter. There is an
angular (82 ), rather narrow internal ledge which reduces
the aperture between collar and bowl to 0.84 oral diameter.
The egg-shaped bowl reaches its greatest diameter of 1.13
oral diameters near 1.0 oral diameter below the rim, the wall
rounding off from the collar convexly to that level. The
lower bowl is full and convex, and is 0.68 oral diameter in
length; the lateral wall tends toward flattening. The short
aboral horn, of about 0.32 oral diameter, is conical (25°) and
blunt at its tree tip.

The wall is thin, hardly exceeding 0.04 oral diameter in
the ledge and elsewhere much thinner. There is a single
layer of large, elongated, subrectangular secondary prisms,
including the ledge, where most species have several layers.
The surface has a distal aggregate of large round fenestrae
and otherwise is provided with rather small, crowded, in-
terpolated subcircular areas. Coccoliths and rhabdoliths are

8

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

frequent. The cavity of the bowl is cut off from that of the
horn by a depressed diaphragm formed by the inner lamina.
Rarely a second, thicker one is found about halfway down
the horn. A convex closing apparatus may be present. The
wall is not particularly dense.

Length, 88u.

There is some difference in wall structure in various indi-
viduals; a few are profusely packed with coccoliths, others
have small prisms, and commonly there are large fenestrae.
The aboral end is generally blunted, but in a few the tip is
sharply pointed.

Codonelhi amphorclla has a bowl shorter, blunter, and
fuller than that of C. recta but not so wide as that of C. rapa.
Its internal ledge is weaker than in these species, and its
collar is more convex than that of recta, which is a cone.

Recorded from nine stations, five in the Atlantic and four
in the Pacific, as follows: one (16) in the Gulf Stream, one
(20) in the Sargasso Sea, three (22, 24, 30) in the Atlantic-
equatorial region, two (81, 95) in the region of South Pacific
island fields, one (139) in the North Pacific trade region,
and one (149) in the California region.

There are 2 pump and 7 net samples, of which 2 were taken
at 50 meters and 7 at 100 meters. Frequencies, minimum.

Temperature: Atlantic, pump sample i9?62, net samples
i5?55-27?88 (20?86); Pacific, 22?78 and 20?4o- 2 8?74
(25^21 ), respectively. Salinity: Atlantic, pump sample 36.48,
net samples 35.61-36.73 (36.14); Pacific, 35.18 and 34.92-
35.85 (35-37), respectively. Density: Atlantic, pump sample
26.01, net samples 23.26-26.34 (24.99); Pacific, 24.15 and
22.4^-24.60 (23.53), respectively. pH: Atlantic, pump sam-
ple 8.17, net samples 7.96-8.30 (8.15); Pacific, 8.28 and 8.19-
8.30 (8.23), respectively.

Codonella apicata Kofoid and Campbell
Codonelhi apicata Kofoid and Campbell, 1929, p. 53, fig. 1 in.
The short, stout, acorn-shaped lorica, with rounded collar,
strong bowl, and faintly pointed aboral end, has a length ol
1.75 oral diameters. The thin-edged oral margin is inturned
and smooth. The rounded collar expands from the margin
to a diameter of 1.15 oral diameters at 0.4 oral diameter
below the rim, and then rapidly contracts to the neck, where
the diameter is 0.95 oral diameter. The sides are full, and
the angle (142 ) at the swollen part is barely rounded off.
The ovate bowl expands from the neck to a diameter of 1.35
oral diameters, which is reached near 0.62 total length below
the rim. Below this level the bowl contracts rapidly to the
decidedly convex-conical (90° to 125 ) aboral region. The
aboral end is bluntly to sharply pointed and less commonly
cuspidate.

The wall averages about 0.05 oral diameter in thickness
across the bowl and is much thinner in the upper collar.
There is only a single layer of large, rectangular secondary
prisms, except in the lower bowl, where two layers are found
in some individuals. There is scarcely any nuchal ledge.
The external surface is reticulated with coarse secondary
polygons with rather thick walls; smaller polygons are sub-
oral, and large "duplex" polygons are common postequa-
torially.

There is a conical closing apparatus.

Length, 52 to 94"..

The length is variable and doubtless reflects the relation
between temperature and available material for lorication.
The wall and the shape of the aboral end are among the
more important variable characters. The "duplex" or ter-
tiary structure is especially interesting.

Codonella apicata is smaller than C. perforata and has a
less angular collar and distinctly different bowl. Codonella
tropica and C. cuspidata have differently shaped collars, and
C. acerca is less trim and also broader. Had it no collar, C.
dadayi would be close to apicata.

Recorded from forty-five stations, thirteen in the Atlantic
and thirty-two in the Pacific, as follows: two (2, 16) in the
Ciulf Stream, one (4) in the Atlantic drift, four (18, 19, 20,
20-21) in the Sargasso Sea, five (22, 23, 24, 25, 27) in the
Atlantic equatorial region, one (31) in the Caribbean Sea,
two (37, 153) in the Pacific equatorial region, six (41, 45,
77, 78, 79, 80) in the Galapagos region, four (57, 63, 65, 67)
in the South Pacific middle latitudes, nine (81, 83, 84, 85, 87,
90, 92, 94, 95) in the region of South Pacific island fields,
four (100, 10 1, 138, 151) in the North Pacific trade region,
five (133, 135, 136, 137, 148) in the California region, and
two (141, 144) in the North Pacific middle latitudes.
Codonella apicata is widely distributed in the warmer regions
and is lacking, save for chance drifting loricae, in cooler
areas. Its absence from stations 6 to 13 and 118 to 125 is
significant. It reaches its maximum near the tropics, where
it may be expected in nearly any sample.

There are 34 pump and 29 net samples, of which 9 were
taken at the surface, 31 at 50 meters, and 23 at 100 meters.
This species apparently prefers deeper levels to the surface,
but is not limited to any special level. Maximum frequency,
20 per cent at station 65; other records above minimum (2 to
7 per cent) from stations 4, 16, 20, 20-21, 22, 45, 63, 77, 78,
80, 81, 85, 136, 148; averages, 14.0 and 7.0 loricae in Atlantic
and Pacific pump samples, and 19.0 and 5.0 per cent in
Atlantic and Pacific net samples, respectively.

Temperature: Atlantic, pump samples i4?32-25?72
(21^74), net samples i4?6o-27?53 (23? 14); Pacific, 14^55-
28?05 (23?38) and i5?03~28?74 (22?52), respectively.
Salinity: Atlantic, pump samples 36.00-37.15 (36.49). net
samples 35.22-38.81 (36.44); Pacific, 31.68-36.49 (35.12) and
34.30-36.98 (35.47). respectively. Density: Atlantic, pump
samples 24.34-26.91 (25.38), net samples 23.84-26.62 (25.15);
Pacific, 20.20-26.11 (23.77) and 22 43 _2 545 ( 2 4-30, respec-
tively. pH: Atlantic, pump samples 7.93-8.32 (8.18), net
samples 8.15-8.27 (8.22); Pacific, 7.92-8.39 (8.27) and 8.01-
8.39 (8.16), respectively.

Codonella aspera Kofoid and Campbell

Codonella aspera Kofoid and Campbell. 1929. pp. 55 - 5 f '- fig-

101.
Codonella galea, Hofker (part), 1931, pp. 352-354, figs. 26, 28

(see also C. galea).
Petalotricha galea, Haeckel, 1899, pi. 3, fig. 6.

The stout, ovoid lorica, with flaring collar, rotund ovate
bowl, and broadly rounded aboral end, has a length of 1.57

TAXONOMY AND DISTRIBUTION

oral diameters. The thin-edged, entire oral rim is surrounded
by a narrow hyaline hand. The collar is an inverted seg-
ment of a cone (35 ) with a length of nearly 0.41 oral diam
etcr, and with a similar diameter at the lower end; the sides
are slightly convex in the middle. The ovate bowl expands
rapidly from the neck, reaches 1.07 oral diameters at its
middle, and then slowly contracts to the blunted, but broad,
subrotund aboral end.

The wall is thickened in the bowl to 0.05 oral diameter,
hut lessens in the fundus and is much thinner in the collar.
There are single, or rarely double, layers ol large, coarse
tertiary structures. The collar is almost transparent, and the
bowl is decidedly dense. The outer surface has large, crudely
formed polygons of various shapes and sizes, and almost
invariably there are large blobs of alveolar material adherent
on the surface, especially in the equatorial region of the bowl.

Length, 85 to 90(1.

Codonella aspera is variable in the shape of the aboral end
and in the extent to which adherent matter is found on the
surface.

Codonella aspera has a more rotund, less trim bowl than
C. elongata and has coarse adherent matter that elongata
lacks; its aboral end is less pointed and its collar has lateral
convexity. Codonella grahami, a close relative of aspera, lacks
surface blobs, has a less rounded bowl, and has less irregular
prismatic structure in the wall. Codonella aspera is a species
of warmer latitudes than grahami, although not tropical.

Recorded from four stations in the Pacific, as follows: one
(128) in the North Pacific middle latitudes, and three (130,
131, 132) in the California region.

There are 2 net and 4 pump samples, of which 4 were
taken at 50 meters and 2 at 100 meters. Maximum fre-
quency, 99 per cent at station 128 at 100 meters; two other
records above minimum (12 and 14 per cent) from same
station at 50 and 100 meters respectively; averages, 50 per
cent and 7 loricae, respectively, in net and pump samples.

Temperature: net samples io?23-i3?98 (i2?io), pump
samples io?23-i8?38 (i3?35). Salinity: net samples 33.14—
33.24 (33.19), pump samples 33.07-33.89 (33.37). Density:
net samples 25.00-25.48 (25.24), pump samples 24.35-2s.4N
(25.37). pH: net samples 8.06-8.39 (8-22), pump samples
8.06-8.33 ( 8 -' 8 )-

Codonella diomedae Kofoid and Campbell
Codonella diomedae Kofoid and Campbell, 1929, p. 59, fig. 118.

The large lorica, with convex, swollen collar, elongated,
saccular bowl, and slightly flattened aboral end, has a length
of 2.17 oral diameters. The oral margin is thin and entire.
There is a low (0.1 oral diameter), hyaline ring around the
opening. The convex, suborally inturned collar expands
rapidly from the rim, and reaches 1.15 oral diameters at 0.33
oral diameter below the rim; the lower two-thirds contracts
to the neck, the diameter of which is equal to that of the oral
opening. The sides are full, and the angle ( 135 ) is clearly
rounded. The long bowl expands below the throat to about
1. 1 5 oral diameters at 0.31 oral diameter below the throat,
becomes 1.23 oral diameters near the equator, and then slowly

contracts in the posterior region to the aboral end. The
aboral end is nearly ll.it (about 0.78 oral diameter in diam-
etei ); commonl) it is jus) rounded ell.

The wall is 0.05 oral diameter in thickness across the bowl
ami thinner in the collar. There are only single layers of
large, rectangular secondary polygons. The external surface
has tine reticulations with small polygons, and rarely "du-
plex" areas.

Length, 105 to 12711.

Codonella diomedae resembles C. galea in general form.
It differs, however, in the swollen bowl and the usual flat
aboral cm\. These characters serve pretty much to distin-
guish it from all the other species.

Recorded from sixteen stations in the I'.i ific, as follows:
four (41, 45, 47, 69) in the Galapagos region, three (62-63,
(>S, ''7 ) in the South Pacific middle latitudes, one (85) in the
region of South Pacific island fields, one (109) in the North
Pacific trade region, four (134, 146. 147, 14.H) in the Cali
fornia region, one (144) in the North Pacific middle l.iti
tudes, and two (152, 153) in the Pacific equatorial region.

There are 6 pump and 11 net samples, of which 4 were
taken at the surface, 3 at 50 meters, and 10 at 100 meters.
Maximum frequency, 6 per cent at stations 65, 67; other
records above minimum (2 to 3 per cent) from stations 109,
144, 147, 152; average in net samples, 2.5 per cent.

Temperature: pump samples i7?4&-22?73 (2i?28), net
samples n?48-27?89 (i9?43). Salinity: pump samples
34.57-36.02 (35.06), net samples 34.19-36.24 (34.92). Den-
sity: pump samples 23.83-25. u (24.46), net samples 23.38-
26.50 (24.79). pH: pump samples 8.12-8.37 (8.24), net
samples 7.76-8.29 (8.13).

Codonella elongata Kofoid and Campbell

(Figure 18)

Codonella elongata Kofoid and Campbell, 1929, pp. 59-60, fig.
102.

The tall lorica, with funnel-shaped collar, long, rather
narrow bowl, and narrowed aboral end, has a length of 1.6
oral diameters. The thin-edged oral margin is entire and
sharp. There is a narrow, hyaline cuff below the margin
with a width of 0.02 oral diameter. The flaring, inverted,
plane, funnel-shaped collar (38 ) has a length of nearly 0.37
oral diameter, and a diameter at the neck of 0.75 oral diam-
eter. Just above the neck there is a narrow (0.07 oral diam-
eter) locally constricted region. The elongated, ovoid bowl
expands from the neck to 1.1 oral diameters at 0.4 its own
length. The sides are somewhat flattened in the equatorial
region, and above and below. The aboral two-fifths is
plainly convex conical (90 ), and the aboral end is narrowly
rounded without distal prolongation.

The wall is 0.05 oral diameter in thickness in the equa-
torial region and gradually thins in the collar. There is a
single layer of large, rectangular prisms. The exterior has
thick-walled secondary polygons, and also in some loricae
an equatorial band of circular tertiary ones of two sizes.

Length. 85 to 1 1711.

The Carnegie loricae are broader-bowled and less trim

10

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

than is typical, and have occasional tertiary wall structure.

Codonella elongata differs from its close relative C. galea
in the more elongated bowl and more nearly pointed aboral
end. The collar is less convex than that of C. aspera, as well
as more regularly contoured and more pointed distally. The
bowl is not so broad as that of C. tropica', and the collar is
not swollen as in the longer C. diomedae.

Recorded from ten stations, seven in the Atlantic and three
in the Pacific, as follows: four (17, 18, 19, 20) in the Sar-
gasso Sea, three (22, 23, 24) in the Atlantic equatorial region,
one (68) in the Galapagos region, one (115) in the North
Pacific middle latitudes, and one (131) in the California
region.

There are 4 pump and 15 net samples, of which 1 was
taken at the surface, 8 at 50 meters, and 10 at 100 meters.
Maximum frequency, 25 per cent at station 115; other records
above minimum (2 to 12 per cent) from stations 18, 19, 20,
22, 24, 131; average in Pacific net samples, 10 per cent; other
averages, 1.2 to 3.6 per cent.

Temperature: Atlantic, pump samples 2o?32-25?72 (23 ?39),
net samples i4?6o-26?63 (20?3o); Pacific, net samples
I2?i2-i6?56 (i4?84). Salinity: Atlantic, pump samples
36.60-37.15 (36.79), net samples 35.61-38.60 (36.11); Pacific,
net samples 33.36-34.85 (34.27). Density: Atlantic, pump
samples 24.34-26.07 (25.22), net samples 24.35-26.62 (25.45);
Pacific, net samples 25.31-25.52 (25.39). pH: Atlantic,
pump samples 8.21-8.27 (8-24), net samples 7.93-8.34 (8.18);
Pacific, net samples 8.10-8.24 (8.15).

Codonella galea Haeckel

(Figures 14, 17)

Codonella galea, Kofoid and Campbell, 1929, p. 60, fig. 106;

Hofker (part), 1931, pp. 352-354, fig. 27 (see also C. aspera).

Not Petalotricha galea, Haeckel, 1899, pi. 3, fig. 6 (see C. aspera).

The moderately tall lorica, with funnel-like collar, gently
constricted throat, ovate bowl, and broadly rounded aboral
end, has a length of 1.41 to 1.57 oral diameters. The thin,
smooth oral margin is sharp-edged. The collar is an in-
verted funnel (25 to 43 ), sometimes with convex sides,
and with a length of 0.32 to 0.38 oral diameter. The diam-
eter at the neck is 0.80 to 0.88 oral diameter. The rather
short, rotund bowl expands evenly from the throat, reaches
a little over 1.0 oral diameter at its middle, and then evenly
rounds off to the broadly rounded to somewhat flattened
aboral end.

The thin wall averages 0.05 oral diameter at the equator
of the bowl and thins to half as much in the collar. Only a
single layer of rectangular prisms occurs (rarely two). The
surface has large polygons with rare larger circular ones
which enclose several of the smaller. The lorica figured
(fig. 14) has a large alveolar blob on the bowl. The collar
is hyaline and the bowl dense.

A conical closing apparatus and sac enclose the whole
body. There are 8 macronuclei, about 12 to 18 mem-
brandies, and a powerful ciliary membrane.

Length, 78 to I20[i.

Like many other widely distributed species, Codonella

galea varies in most characters, as a survey of the literature
clearly shows. For this reason several names have been
applied to galea (for summary see Kofoid and Campbell,
1929). Most of Hofker's ( 1931 ) material belongs to C.
aspera, but his figure 27 is of galea.

Codonella galea differs from C. nationahs in being taller,
with a less stout bowl and a more flaring collar. Other re-
lated species, e.g. C. elongata, tropica, cuspidata, robusta, and
aspera, differ mainly in the shape of the aboral region or
have different collars.

Recorded from thirty-three stations, twelve in the Atlantic
and twenty-one in the Pacific, as follows: two ( 14, 16) in the
Gulf Stream, three (18, 19, 20) in the Sargasso Sea, six (22,
23, 25, 26, 27, 28) in the Atlantic equatorial region, one (33)
in the Caribbean Sea, six (40, 45, 46, 71, 77, 78) in the
Galapagos region, two (54, 65) in the South Pacific middle
latitudes, two (82, 85) in the region of South Pacific island
fields, three (100, 140, 151) in the North Pacific trade region,
five (130, 135, 137, 147, 148) in the California region, two
(142, 145) in the North Pacific middle latitudes, and one
(153) in the Pacific equatorial region.

There are 21 pump and 27 net samples, of which 5 were
taken at the surface, 21 at 50 meters, and 22 at 100 meters.
This species appears to be subsurface by preference. Maxi-
mum frequency, 6 per cent at stations 137, 147; other
records above minimum (2 to 5 per cent) from stations 14,
16, 18, 23, 25, 26, 45, 46, 54, 77, 130, 145, 148, 151, 153;
average in Pacific net samples, 3.3 per cent; other averages,
1.6 to 2.2 per cent.

Temperature: Atlantic, pump samples ^95-24? 10 (20?2i),
net samples 1 4^60-26? 04 (2i?68); Pacific, i6?96-27?62
(23?i7) and i2?9i-24?84 (2o?5g), respectively. Salinity:
Atlantic, pump samples 35.10-36.81 (36.01), net samples
35.70-38.18 (36.33); Pacific, 33.70-36.46 (35.34) and 33.40-
36.04 (34.92), respectively. Density: Atlantic, pump samples
24.49-26.08 (25.44), net samples 23.98-26.62 (25.30); Pacific,
22.31-25.14 (24.56) and 23.50-25.37 (24.52), respectively.
pH: Atlantic, pump samples 8.18-8.22 (8.21), net samples
7.93-8.27 (8.19); Pacific, 8:10-8.34 ( 8 - 22 ) and 8.12-8.34
(8.22), respectively.

Codonella grahami, new species
(Plate 1, figure 6)
The lorica is moderately elongated, and pointed. It has a
length of 2.06 oral diameters. The oral rim is smooth and
slightly rounded. The collar is the basal segment of an in-
verted truncated cone (27°); its length is 0.52 oral diameter,
and the diameter at the aboral end, which forms the con-
stricted neck, is 0.87 oral diameter. The sides of the collar
are regular and smooth and there is little, if any, median
bulge. The bowl as a whole is elongated, and olive-like in
shape. Its suboral part, which joins with the neck, is a
rounded segment of a cone (about 45°), and the basal sec-
tion is the level of the greatest diameter of the bowl, 1.13
oral diameters, reached near 0.45 total length from the rim.
The lower part of the bowl is an inverted cone (53°) with
full, curved sides. The aboral end is pointed but not pro-

TAXONOMY AND DISTRIBUTION

II

longed or pedicellated. The sides ol the lorica lorm, in side
view, a sigmoid flexure with the lower arm prolonged.

The wall is subuniform in thickness at all levels and
approximately o.oi oral diameter in thickness. There is a
slight thickening at the neck. The wall is coarsely and
irregularly prismatic, with large, ovoid tertiary structure and
enclosed secondary prisms. The tertiary prisms are largest at
the ambitus of the howl and smaller elsewhere-, especially in
the collar; the thin oral rim is hyaline. The lumen, except
for the nuchal thickening, follows the outer contour.

Length, total Sy\l, collar l8ii; diameter, oral J7(i, maxi-
mum 530..

Codonella grahami is rather uniform in its general char-
acters, and the many loricae examined were much alike. The
surface meshwork, however, is very variable in the sizes and
shapes of the prisms; these may be hexagons, pentagons, or
subcircles.

Codonella grahami resembles C. clongata in general form,
but the aboral end is distinctly pointed instead ot evenly
rounded. It never has coccoliths, which clongata often has.
It bears some likeness to C. aspera, but that species has a
more rounded bowl, a rotund aboral end, and a coarser and
more irregular prismatic structure; it often includes blobs of
alveolar material in its walls, and grahami consistently lacks
these. The latter is much narrower and thinner than C.
acuta, although both species are aborally pointed; the bowl
of acuta reaches its maximum at a lower level than does that
of grahami, and it frequently has many large coccoliths,
which are lacking in grahami. Codonella grahami also
comes from generally cooler waters than do most others of
the genus.

Recorded from five stations in the Pacific, as follows: four
(in, 115, 116, 117) in the North Pacific middle latitudes,
and one (118) in the East Asiatic marginal sea.

There are 5 pump and 2 net samples, of which 1 was taken
at the surface, 4 at 50 meters, and 2 at 100 meters. Maxi-
mum frequency, 6 per cent at stations in, 117; one other
record above minimum (4 per cent), from station 117; re-
mainder at minimum; averages, 5 per cent and 2 loricae in
net and pump samples, respectively.

Temperature: net samples 8?93~i5?56 (i2?74), pump
samples g°jy-ig°.^() (13:55). Salinity: net samples 34.06-
34.22 (34.14), pump samples 33.61-34.63 (34.23). Density:
net samples 25.89-26.41 (26.15), P um P samples 24.62-26.27
(25.62). pH: net samples 7.98-8.06 (8.02), pump samples
8.02-8.21 (8.09).

Type locality, station 117, at 100 meters; latitude 40 20'
north, longitude 150 58' east.

Codonella nationalis Brandt

Codonella nationalis, Kofoid and Campbell, 1929, p. 63, fig.
107; Hofker, 1931, pp. 356-357, fig. 30.

The short, rather stout, potlike lorica, with fairly regular
rim, rotund collar, squat bowl, and broad aboral end, has a
length of 1.35 oral diameters. The oral margin is thin-
edged, minutely but highly regularly denticulate, and located
above a variable hyaline band with a width ol approximately
0.03 oral diameter. The collar is a segment of an inverted

funnel (within io°), and the diameter at the neck is 0.89
oral diameter. A bare nuchal ledge is present. The plump
bowl expands from the neck and reaches its maximum diam-
eter of 1.08 oral diameters at the laterally flattened equator.
The bowl gently contracts, reaching O.93 oral diameter at
o.S> total length below the rim, and then rapidly rounds oil
aborally.

The wall averages 0.04 oral diameter in thickness in the
bowl and thins out 111 the collar. There are two layers of
large, irregular, rectangular to pentagonal secondary poly-
gons; primary alveoles are enclosed by them. The external
surface has a network of coarse polygons and enclosed
smaller ones.

Length, 77 to 1 171A.

Codonella nationalis resembles C. inflata but is more trim,
with greater nuchal constriction, less wide bowl, and Hatter
aboral end. Codonella tropica and C. cuspidata have pointed
aboral ends, as does also the much smaller C. acerca.

Recorded from ten stations, five each in the Atlantic and
Pacific, as follows: four (17, 18, 19, 20) in the Sargasso Sea,
one (28) in the Atlantic equatorial region, two (78, 80) in
the Galapagos region, two (81, 85) in the region of South
Pacific island fields, and one (150) in the North Pacific trade
region.

Net samples only, 1 1 in all, 6 taken at 50 meters and 5 at
100 meters. Maximum frequency, 7 per cent at station 81;
other records above minimum (2 to 5 per cent) from stations
17, 18, 19, 78, 80, 85, 150; averages, 2.2 and 4.4 per cent,
the latter in the Pacific.

Temperature: Atlantic, i9?82-25?3i (22?58); Pacific,
i9?27-27?89 (24?8o). Salinity: Atlantic, 36.60-37.15 (36.83);
Pacific, 34.63-36.24 (35.74). Density: Atlantic, 24.89-26.05
(25.44); Pacific, 23.38-24.68 (23.94). pH: Atlantic, 8.19—
8.27 (8.23); Pacific, 8.14-8.32 (8.21).

Codonella olla Kofoid and Campbell
(Figures 15, 16)
Codonella olla Kofoid and Campbell, 1929, pp. 63-64, fig. 115.

The decidedly potlike, squat lorica, with widely expanded
and flattened aboral end, has a length of 1.4 oral diameters.
The oral margin is coarsely undulating and minutely irregu-
lar, thin, and inturned. The collar bulges; its length is a
trifle less than 0.33 oral diameter, with the same diameter at
the throat as at the upper end, but the bulge in the middle
is 1.1 oral diameters. There is a scarcely developed, angular
(8o°) internal ledge at the throat, which reduces the diam-
eter of the opening there to 0.83 oral diameter. The bowl
swells from the neck to a diameter of 1.33 oral diameters at
0.67 total length from the rim. Below this level, it gradually
contracts to the nearly flat aboral end, the diameter of which
is 0.67 oral diameter.

The thin wall (which reaches, however, nearly 0.09 oral
diameter in thickness across the ledge) is not over 0.02 oral
diameter in the bowl. There are single to triple layers of
subrectangular, hyaline secondary prisms. Over the whole
surface are large, subcircular, clear tenestrae between which
are rather faint smaller circles or alveoles.

12

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Length, 75 to 90^,.

One of the loricae figured (fig. 16) is unusual. Its length
is 1.53 oral diameters. Its howl is distinctly elongated and
reaches its greatest diameter at little above the flat distal
end, and its whole surface is studded with elongated rhab-
doliths. It is go|x in length. The oral rim is entire.

Codonella olla resembles C. poculum closely, but differs in
the expanded and flattened aboral end. Its internal shelf is
less developed, there is constriction at the throat, and its
collar bulges. Its aboral end is flat, unlike that of C. acerca,
which is pointed.

Recorded from three stations in the Atlantic, as follows:
two (19, 20) in the Sargasso Sea, and one (27) in the
Atlantic equatorial region.

Net samples only, of which 1 was taken at 50 meters and
2 at 100 meters. Maximum frequency, 2 per cent at station

Temperature, 22?42-26?04 (>3?7o); salinity. 36.25-37.03
(36.71); density, 23.98-25.67 (25.01); pH, 8.19-8.30 (8.24).

Codonella poculum Kofoid and Campbell
Codonella poculum Kofoid and Campbell, 1929, p. 64, fig. 114.
The distinctly stout, pot-shaped lorica, with collar undif-
ferentiated except by an internal ledge and with rounded
bowl, has a length of 1.1 oral diameters. The oral margin
is regularly denticulate, there being approximately 50 low,
wide, equidistant, regular, triangular teeth. These hyaline
teeth arise from a narrow, similar band, the width of which
is less than 0.2 oral diameter. The collar is distinguished
only by the internal ledge or shelf located near 0.3 oral diam-
eter below the rim. The collar is virtually a cylinder with
only minor, strictly local modifications in external contour.
The internal shelf is triangular (75 ) and, because of its
width, reduces the opening between collar and bowl to 0.73
oral diameter; its base, against the outer wall, is 0.16 oral
diameter in width. The bowl continues the subcylindrical
form of the collar for 0.71 total length with a few local ir-
regularities, and has a diameter at this level of 0.95 oral
diameter. The aboral region rapidly rounds off so that the
thimble-shaped end is 0.29 total length below the level of the
cylindrical upper section.

The relatively thick wall reaches, in the mid-region of the
bowl, as much as 0.05 oral diameter; in the collar it is about
half as much. There are one to five layers of faint, rounded
to rectangular secondary prisms as well as a lesser number
of much larger tertiary ones. The outer surface shows these
prisms in circular shapes.

There is a strong closing apparatus.
Length, 80 to 92|i.

Codonella poculum closely resembles C. acutula, but its
aboral end is rounded instead of pointed as in acutula. It
lacks the nuchal constriction which is found in C. olla. The
aboral end is less flattened, not expanded, and the wall is
different in these two species. Codonella acerca, aside from
its smaller size, has a pointed aboral end and also nuchal
constriction.

Recorded from four stations in the Pacific, as follows: one

(99) in the Pacific equatorial region, two (101, 150) in the
North Pacific trade region, and one (148) in the California
region.

Pump samples only of which 1 was taken at the surface,
1 at 50 meters, and 2 at 100 meters. Frequency, minimum.

Temperature, i9?27~27?93 (23?38); salinity, 34.63-34.94
(34.81); density, 22.39-24.72 (24.14); pH, 8.21-8.32 (8.25).

Codonella rapa Kofoid and Campbell
(Figure 10)
Codonella rapa Kofoid and Campbell, 1929, p. 65, fig. 130.
The short, stout, deep-throated lorica, with swollen collar,
plump bowl, and small conical aboral horn, has a length of
1.74 oral diameters. The minutely subdenticulate, rather
regular oral margin is thin-edged. The short, bulging collar
has a length of 0.43 oral diameter; its diameter across the
middle is 1.09, and at the throat 0.95 oral diameter. Inside
the throat is an angular (8o°), rather wide shelf that reduces
the aperture to 0.73 the external diameter. The swollen
bowl is rotund, reaches its greatest diameter of 1.1 oral
diameters near 1.0 oral diameter below the oral rim, and
then rather gradually narrows down to the aboral end, where
there is a conical (34°) aboral horn (0.26 oral diameter in
length) with widely flaring (8o°) base and sharply pointed
free tip.

The wall is thickest in the upper bowl, where it may attain
0.08 oral diameter, is reduced gradually in the collar and
fundus to less than 0.02 oral diameter, and contains, mostly,
only a single layer of large, rectangular prisms. The sur-
face shows small hexagonal to circular prisms (primary?),
and often each of the large secondary areas has a large cocco-
lith enclosed in it. The aboral horn is hollow, but its cavity
is cut off from that of the bowl by a thin-walled diaphragm
formed of the inner lamina.
Length, 781X.

The Carnegie loricae are not so stout as those described by
Kofoid and Campbell (1929).

Codonella rapa has a more flaring collar, shorter bowl,
and stouter horn than C. recta. Codonella amphorella has a
longer bowl with less rotundity. These three attractive
species of the high latitudes can scarcely be confused with
others of the tropical oceans.

Recorded from two stations (62-63, 64) in the South
Pacific middle latitudes.

Net samples only, 1 taken at the surface and 1 at 1000
meters. There were 2 loricae.

Temperature (at 1000 meters) 3^98; salinity, 34.30; den-
sity, 27.25; pH, 7.76.

Codonella recta Kofoid and Campbell
Codonella recta Kofoid and Campbell, 1929, p. 65, fig. 131.
The stout lorica, with shallow throat, erect collar, and
stout horn, has a length of 1.78 oral diameters. The oral
margin is very thin, erect, and entire. The collar is sub-
conical (io°) with a length of 0.4 oral diameter; its lower
diameter is only a little less than the oral diameter, and the

TAXONOMY AND DISTRIBUTION

13

slightly developed, angular (70 ) internal ledge reduces the
aperture between collar and bowl to 0.87 oral diameter. Its
sides are practically plane except that they rather suddenly
bend inward a little just above the throat. The ellipsoidal
bowl has a maximum diameter of 1.1 oral diameters slightly
below its middle. Below this level it gradually contracts to
the aboral horn. The horn has a wide, asymmetrical conical
(50 ) base and narrower conical (io°) stem. Its free tip
is blunted.

The wall is very thin, being not over 0.02 oral diameter in
thickness in the bowl. There is a single or double layer of
narrow, subrectangular to irregular secondary prisms, within
which are closely packed primary alveoles. The collar has
large subcircular fenestrae which gradually merge in the
bowl with smaller circles. These circular structures have
thick walls in which minute alveoles are present. The
lumen of the horn is cut off from that of the bowl by a
depressed diaphragm. The wall is dense.

Length, 951.1.

Codonella recta has a conical collar which lacks the bulge
of that of C. amphorella or C. rapa. The bowl is less rotund
and there is much less constriction at the throat than in
either of the above two species.

Recorded from one station (154) in the Pacific equatorial
region, in a net sample taken at 100 meters. Frequency, 2
per cent.

Temperature, 25?8i; salinity, 34.82; density, 22.98; pH,

7-93-

Codonella tropica Kofoid and Campbell
Codonella tropica Kofoid and Campbell, 1929, p. 67, fig. no.

The stout, ovate lorica, with bulging collar, wide bowl,
and simple, pointed aboral end, has a length of 1.34 oral
diameters. The thin-edged oral margin is entire, and below
it is a narrow, hyaline cuff, the width of which is about 0.1
oral diameter or less. The convex collar expands from the
rim to 1.08 oral diameters near its middle and then some-
what more rapidly contracts to 0.92 oral diameter at the
neck. The collar has a length of nearly 0.42 oral diameter.
The stout, short bowl expands from the throat and reaches
1. 16 oral diameters at 0.66 oral diameter below the rim.
Below this widest level it rapidly contracts, with full, convex
sides, to the pointed end, which bears a minute, extended
point.

The wall reaches approximately 0.05 oral diameter in
thickness in the bowl and is thinner in the collar. There is
a single layer of large, rectangular prisms. The exterior
meshwork is made up of small, subcircular polygons, every-
where of uniform size.

Length, 7711.

Codonella tropica is stouter, with less constricted throat,
shorter bowl, and more pointed end than C. acuta. The wall
of the latter also bears large coccoliths. Codonella cuspidata
is close to tropica, but less wide and hence more slender; it
has a larger collar with greater nuchal constriction.

Recorded from twelve stations in the Pacific, as follows:
one (77) in the Galapagos region, four (81, 82, 84, 95) in
the region of South Pacific island fields, two (112, 113) in

the North Pacific middle latitudes, one (151) in the North
Pacific trade region, three (152, 153, 154) in the Pacific
equatorial region, and one (146) in the California region.

There were 2 pump and n net samples, of which 1 was
taken at the surface, 5 at 50 meters, and 7 at 100 meters.
Maximum frequency, 6 per cent at stations 153, 154; other
records above minimum (2 to 3 per cent) from stations 81,
146, 152.

Temperature: pump samples 20?i6-23?77 (2i?96), net
samples ii?48-28?74 (22^92). Salinity: pump samples
34.59-34.71 (34.65), net samples 34.32-36.42 (35.22). Den-
sity: pump samples 23.42-24.52 (23.97), net samples 22.43-
26.50 (24.06). pH: pump samples 8.20-8.25 (8-22), net
samples 7.76-8.28 (8.14).

CODONARIA Kofoid and Campbell
Codonaria Kofoid and Campbell, 1939, p. 55.

The genus Codonaria includes a number of species from
the warmer parts of the sea. They have clearly been derived
from Codonella. Nearly all are inhabitants of the open
ocean or of the Mediterranean. They are rare in the South-
ern Hemisphere save near the equator.

Six species are described.

Codonaria angusta Kofoid and Campbell
Codonella angusta Kofoid and Campbell, 1929, p. 53, fig. 1 17.

The rotund lorica, with tall, convex collar, deep throat,
and distally inflated bowl, has a length of 1.56 oral diam-
eters. The thin, smooth oral margin arises from a hyaline
cuff only 0.04 oral diameter in width. The inflated collar is
subcylindrical, with a length of 0.32 total length, with its
greatest diameter of i.iforal diameters at 0.67 of its length
from the upper end, and with the barely developed flangelike
ledge at 0.18 oral diameter below the rim; this narrow ledge
scarcely projects beyond the collar, and is sharply angular
(32 ). The constricted neck of the bowl has a diameter of
0.98 oral diameter. The rotund bowl expands rapidly from
the throat, reaches 1.21 oral diameters near 0.6 total length
from the rim, and then contracts sharply to the nearly flat-
tened aboral end (0.65 oral diameter), which lacks any trace
of special differentiation.

The wall of the bowl is nearly 0.06 oral diameter in thick-
ness, that of the internal ledge is twice as much, and in the
collar the wall rapidly declines to one-tenth as much. There
are one to three layers of thick-walled, irregular, elongate,
alveole-like secondary prisms. The truncate, subtriangular
internal ledge at the junction of collar and bowl reduces the
cavity to 0.83 the external diameter. The exterior surface
has large and small, rather faint subpentagonal to subhex-
agonal meshes.

Length, 80 to 98^.

Codonaria angusta may be distinguished by the narrow
external and thick internal ledges, the laterally convex collar,
and the swollen, flattened bowl. In a few respects it is
transitional between Codonella diomedae and Codonaria
cistellula.

Recorded from one station (19) in the Sargasso Sea,

14

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

in a net sample taken at 50 meters. Frequency, minimum.
Temperature, 25?3i; salinity, 37.15; density, 24.89; pH,
8.27.

Codonaria benguelensis Kofoid and Campbell
Codonclla benguelensis Kofoid and Campbell, 1929, p. 57, fig.

111.

The stout, potlike lorica, with tall suboral cone and collar,
and plump bowl, has a length of 1.56 oral diameters. The
very thin, irregular oral margin has a cuff about 0.12 oral
diameter in length below it. The suboral cone (50 ) with
laterally concave sides has a length of nearly 0.15 oral diam-
eter, and a diameter of 1.1 oral diameters at its lower end.
The suboral ledge is separated from it by an angular (60°)
trough. The thick, slightly wavy, shelflike ledge has a
diameter of 1.22 oral diameters, and its thickness reaches
0.07 oral diameter. The collar is an inverted basal segment
of a truncated cone (17°) with a length of nearly 0.5 oral
diameter, and a diameter of a trifle more than 1.0 oral
diameter at its lower end. The suboral cone and the collar
together give the upper region of the lorica a decidedly con-
vex form. The rotund, rather squat bowl reaches its greatest
diameter of 1.17 oral diameters at 1.0 oral diameter below
the rim, below which level it gradually rounds off to the
blunt, subhemispherical aboral end.

The wall is thickest in the collar and bowl, where it is less
than 0.06 oral diameter. A single layer of large, thick-
walled, rectangular secondary prisms is everywhere present
save in the internal ledge, where there are two layers. The
cuff is hyaline, the suboral cone and collar and the ledge
have moderately large, thick-walled hexagons, and the bowl
has very large "duplex" circular fenestrae in a wide equa-
torial band. The internal angular ledge at the junction of
collar and bowl reduces the aperture between them to 0.85
oral diameter.

Length, 80 to 88u\.

Codonaria angusta has a more convex suboral section less
sharply distinguishable into cone and collar, and also a
thinner, less evident ledge than does C. benguelensis. The
former, too, lacks duplex structure, and its bowl is not par-
ticularly similar. Codonaria oceanica has a wider bowl and
lower suboral region.

Recorded from one station (100) in the North Pacific
trade region, in a net sample taken at 50 meters. Only 1
lorica was present.

Temperature, 1-]%"]; salinity, 34.71; density, 22.31; pH,
8.22.

Codonaria cistellula (Fol) Kofoid and Campbell

Codonclla cistellula, Kofoid and Campbell, 1929, p. 57, fig. 125;
Hofker, 1931, pp. 354-356, fig. 29.

The rather elongate, tall lorica, with flaring collar, mod-
estly inflated bowl, and pointed aboral end, has a length of
2.0 oral diameters. The minutely subdenticulate oral margin
is thin and there is a low cuff below the rim. The suboral
cone is a slightly concave basal segment of a cone (25 ) with
a length of only 0.11 oral diameter, and with a basal diam-
eter of 1.26 oral diameters. It is separated from the upper

edge of the collar by a shallow, subangular trough. The
ledge is the smooth, free upper edge of the collar, the diam-
eter of which is 1.39 oral diameters. The collar is an in-
verted truncated basal segment of a plane to barely convex
cone (45°) with a length of 0.62 oral diameter and a
diameter at the neck of 1.0 oral diameter. The bowl rounds
off and reaches its greatest diameter at 0.65 total length
below the margin; this maximum is 1.57 oral diameters
(only a little less than the length of the whole lorica). The
wall quickly and convexly rounds to the sharp but not pro-

iged aboral end, which

oral diameter below the level

of the maximum diameter.

The wall is thickest in the lower collar and upper bowl,
where it reaches almost 0.09 oral diameter. It thins out in
the suboral cone and toward the aboral end to about 0.06.
There are one to four, mostly two to three, layers of sub-
hexagonal, thick-walled secondary prisms. The walls of
these prisms, under the best magnification, show primary
alveoles in one or two layers. The cuff is hyaline, but the
remainder of the surface has large, fairly heavy subrectangu-
lar to subhexagonal meshes. Large "simplex" fenestrae are
scattered over collar and bowl, mostly around the equator.
There is a conical closing apparatus of 12 truncated tri-
angular blades and a somatic enclosing sac. There are 12 or
more membranelles, and 16 macronuclei.

Length, 90 to 125^.

Codonaria cistellula is taller and longer than the other
species. Its bowl is less inflated, its aboral end less sharply
pointed, and its maximum inflation lower than in others.
It is not easily confused with them, the aboral characters in
general serving to distinguish it. Had it no suboral cone it
might be confused with Codonella diomedae, which, how-
ever, is a different form.

Recorded from eight stations, three in the Atlantic and
five in the Pacific, as follows: two (17, 18) in the Sargasso
Sea, one (22) in the Atlantic equatorial region, two (69, 75)
in the Galapagos region, two (in, 145) in the North Pacific-
middle latitudes, and one (146) in the California region.

There are 2 pump and 6 net samples, of which 1 was taken
at the surface, 3 at 50 meters, and 4 at 100 meters. Fre-
quency above minimum (2 to 3 per cent) at stations 18, 69,
145, 146.

Temperature: Atlantic, net samples i7?50-2i?85 (i9?89);
Pacific, pump samples i9?39-2i?i3 (20?26), net samples
i8?40-20?07 (i9?2i). Salinity: Atlantic, net samples 36.60-
36.81 (36.70); Pacific, pump samples 34.58-35.24 (34.01).
net samples 34.32-35.47 (34.70). Density: Atlantic, net
samples 25.49-26.07 (25.78); Pacific, pump samples 24.62-
24.66 (24.64), net samples 24.24-25.55 (24.89). pH: Atlan-
tic, net samples 8.21-8.27 (8.24); Pacific, 1 pump sample
8.12, net samples 8.10-8.34 (8.23).

Codonaria lata Kofoid and Campbell

(Figure 12)

Codondla lata Kofoid and Campbell, 1929, p. 62, fig. 126.

The stout, small lorica, with flaring collar, rotund bowl,
and flattened aboral region, has a length of 1.47 oral diam-

TAXONOMY AND DISTRIBUTION

15

eters. The thin oral margin is subdenticulate and enclosed
In an exceedingly low (0.11 oral diameter), suhcylindrical
suboral cuff. The suboral ledge has a diameter of 1.13 oral
diameters and is separated from the cuff by a concave-
angular trough. The rather long, inverted-conical ( 34°)
collar, with plane sides, has a length of little more than 0.4
oral diameter, and a diameter at the constricted throat of
over 0.87 oral diameter. The subglobose bowl reaches its
greatest diameter of r.13 oral diameters near 1.0 oral diam-
eter below the rim; from that level to the aboral end is about
0.53 oral diameter. The aboral end is flattened and has a
diameter of nearly 0.5 oral diameter.

The wall has a thickness of nearly 0.05 oral diameter ex-
cept in the cuff, where it is less than half as much. There
are one to three layers of irregular secondary prisms. The
outer surface has small subcircular reticulations which crowd
in the ledge and near the aboral end. There is an equa-
torial and southern temperate band of subequal circular
fenestrae around the bowl.
Length, 78 to 90U.

The Carnegie loricae tend toward aboral flattening, flat
sides, and steeply pitched flare in the collar.

Codonaria lata is longer than C. australis and has a flatter
aboral end. Its bowl is not so long as that of C. bengue-
lensis or so wide as that of C. oceanica. There is no trace of
the aboral point which distinguishes C. mucronata, dadayi,
or cistellula. The collar lacks the submedian bulge that
characterizes C. angusta.

Recorded from six stations, two in the Atlantic and four
in the Pacific, as follows: one (19) in the Sargasso Sea, one
(28) in the Atlantic equatorial region, one (78) in the Gala-
pagos region, one (113) in the North Pacific middle lati-
tudes, one (109) in the North Pacific trade region, and one
( 146) in the California region.

There are 7 net samples, of which 3 were taken at 50
meters and 4 at 100 meters. Maximum frequency, 8 per cent
at station 113; other records above minimum (2 to 3 per
cent) from stations 78, 109; average in the Pacific, 3.2 per
cent.

Temperature: Atlantic, 25?3i-27?57 (26?44); Pacific,
i9?8i-24?38 (2i?92). Salinity: Atlantic, 36.24-37.15 (36.69);
Pacific, 34.32-36.03 (34.92). Density: Atlantic, 23.49-24.89
(24.19); Pacific, 23.74-24.72 (24.21). pH: Atlantic, 8.27-
8.30 (8.28); Pacific, 8.14-8.26 (8.20).

Codonaria mucronata Kofoid and Campbell

(Figure 13)

Codonella mucronata Kofoid and Campbell, 1929, pp. 62-63,
fig. 123.

The stout lorica, with deep constriction at the throat and
with pointed aboral end, has a length of 1.72 oral diameters.
The thin, subdenticulate oral margin is enclosed by the low
suboral cuff. The cuff is a low band or basal segment of a
concave cone (12 ) with a length of less than 0.1 oral diam-
eter. The cuff is superimposed on the wider, flaring, plane,
irregularly swollen, inverted-conical (58 ) collar, the length
of which is a little over 0.4 oral diameter; its lower diameter

is approximately 0.93 oral diameter and its diameter at the
upper end is 1.34 oral diameters. The upper margin is
roughly undulating although without teeth, and connects
with the suboral cuff by means of a concave-angular trough.
The rather short, squat bowl expands from the constricted
throat, forming a segment of a cone (32 ), and reaches a
diameter equal to that of the upper end oi the collar at 0.67
total length below the rim. Below this widest level the bowl
rapidly contracts as an inverted decidedly convex cone (98 );
the convexity reaches its greatest width within the upper
three-tenths and quickly declines so that the aboral end is
pointed, without, however, being produced.

The wall is uniformly about 0.06 oral diameter in thick-
ness in the collar, and less in the cull There are one to
three layers of irregularly arranged, subrectangular secondary
prisms enclosed within thin laminae.
Length, 75 to io8p:.

The Carnegie loricae are a little longer than those earlier
recorded.

Codonaria mucronata agrees with C. dadayi in the sharply
pointed aboral end, but differs in its wide, more squat bowl,
constricted throat, and better-developed ledge. Codonaria
cistellula, also pointed, differs in being narrower and taller,
and in its fuller ledge. The other species are not likely to
be confused with mucronata, which is easily distinguished
even in rapidly examined plankton.

Recorded from nine stations, four in the Atlantic and five
in the Pacific, as follows: one (14) in the Gulf Stream, two
(17, 18) in the Sargasso Sea, one (33) in the Caribbean Sea,
two (45, 71) in the Galapagos region, and three (152, 153,
154) in the Pacific equatorial region.

There are 4 pump and 10 net samples, of which 2 were
taken at the surface, 4 at 50 meters, 8 at 100 meters. Maxi-
mum frequency, 12 per cent at station 71; other records above
minimum from stations 45, 71, 152, 153, 154; average in
Pacific net samples, 5.1 per cent.

Temperature: Atlantic, net samples i9?82-23?i7 (2^29),
pump sample 14^95; Pacific, ii?48-25?8i (21^24) and
n?48-23?46 (19? 10), respectively. Salinity: Atlantic, net
samples 36.49-36.81 (36.62), pump sample 35.10; Pacific,
34.73-35.24 (35.00) and 34.73-35.24 (35.06), respectively.
Density: Atlantic, net samples 25.03-26.07 (25.66), pump
sample 26.08; Pacific, 22.98-26.50 (24.34) and 24.00-26.50
(24.93), respectively. pH: Atlantic, net samples 8.18-8.27
(8.22), pump sample 8.18; Pacific, 7.76-8.28 (8.05) and
7.76-8.13 (8.00), respectively.

Codonaria oceanica (Brandt) Kofoid and Campbell

Codonella oceanica, Kofoid and Campbell, ^29, p. 63, fig. 122.

The short, wide lorica, with inflated bowl and rather low
collar, has a length of 1.7 oral diameters. The ragged, ir-
regular oral margin is thin-edged and a low cuff lies below
it. The suboral cone (8o°) is a low, truncated, concave
basal segment with a length of nearly 0.17 oral diameter and
a basal diameter of 1.2 oral diameters. The suboral ledge is
not wide (1.3 oral diameters), is widely angular (90 ), and
is not especially differentiated from the upper edge of the

i6

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

collar, of which it forms the wavy, free end. The wide,
plane collar is an inverted truncated cone (50°), the lower
diameter of which is 0.92 oral diameter, reached at a little
over 0.54 oral diameter below the rim. The swollen, wide
bowl attains its greatest diameter of 1.35 oral diameters near
0.66 total length below the rim. The aboral end is widely
but bluntly angular (133°) and undifferentiated.

The wall is thickest in the lower collar and upper bowl,
where it reaches 0.07 oral diameter. It thins above and
below to one-fourth this thickness, or even less. Single
layers of large, rectangular secondary prisms occur except in
the thicker regions, where there are two or even three layers
of smaller, commonly hexagonal ones. The cuff is hyaline,
the suboral cone has small prisms, and the collar and upper
bowl have larger, variously shaped ones, there being as
many as 72 around the equator of the bowl. An internal
ledge is lacking.

Length, 70 to 951-1.

Codonarla oceanica has a wide bowl like C. mucronata,
from which it differs in having a bluntly hemispherical in-
stead of pointed end. The lorica is more squat than that
of C. cistellula, which also is taller. It is not likely to be
confused with the other species.

Recorded from twenty-one stations, eight in the Atlantic
and thirteen in the Pacific, as follows: two (2, 15) in the Gulf
Stream, three (18, 19, 20) in the Sargasso Sea, two (23, 24)
in the Atlantic equatorial region, one (34) in the Caribbean
Sea, one (35) in the Pacific equatorial region, five (40, 45,
46, 71, 78) in the Galapagos region, two (66, 67) in the South
Pacific middle latitudes, one (81) in the region of South
Pacific island fields, two (112, 145) in the North Pacific
middle latitudes, one (136) in the California region, and one
(151) in the North Pacific trade region.

There are 8 pump and 16 net samples, of which 4 were
taken at the surface, n at 50 meters, and 9 at 100 meters.
Maximum frequency, 5 per cent at station 45; other records
above minimum (2 per cent) from stations 2, 34, 46; average
in Pacific net samples, 2 per cent.

Temperature: Atlantic, pumpsamples20?32-24?8i (22?4i),
net samples 20?35~25?3i (22?8i); Pacific, i6?58-23?25
(18764) and i4?33~26?42 (2i?i7), respectively. Salinity:
Atlantic, pump samples 36.39-36.82 (36.67), net samples
36.02-37.15 (36.46); Pacific, 34.60-35.13 (34.87) and 34.42-
36.03 (35.32), respectively. Density: Atlantic, pump samples
24.47-26.07 (25.70), net samples 24.67-25.76 (25.15); Pacific,
23.58-26.21 (24.87) and 24.00-26.06 (24.56), respectively.
pH: Atlantic, pump samples 8.21-8.24 ($-23), net samples
8.14-8.27 (8.19); Pacific, 7.85-8.39 (8.17) and 7.88-8.19
(8.10), respectively.

CODONOPSIS Kofoid and Campbell

Codonopsis Kofoid and Campbell, 1939, pp. 60-61.

Only a single species of this genus is known. It is peculiar
to the western part of the Pacific, not having been reported
elsewhere, even in the Indian Ocean. Evidently the distri-
bution of Tintinnoina is not conditioned solely by tempera-
ture or other physical factors, but by geographical conditions

as well. One other genus of some importance, Epicranella,
is exclusively Pacific, being limited to the cool waters of the
Humboldt Current. No species of Epicranella were found
in Carnegie material, but the Humboldt Current was not
extensively explored. In the Albatross and Zaca material
several species have been found; these two ships have in-
tensively explored the coasts of Peru and Chile.

Codonopsis ollula (Brandt) Kofoid and Campbell

Cyttarocylis ollula, Kofoid and Campbell, 1929, p. 115, fig. 212.

Codonopsis ollula, Kofoid and Campbell, 1939, pp. 61-62.

The stout, egg-shaped lorica, with distinct external sub-
oral ledge below the squarely truncated oral rim, and evenly
blunted aboral end, has a length of 1.41 oral diameters. The
thin, erect oral rim is entirely free of irregularity. The
collar region extends to the external ledge at 0.19 oral diam-
eter below the rim. It is ringlike, and triangular in section,
the apex forming the oral margin. The ledge is a hori-
zontal shelf 1.26 oral diameters in diameter, the sides of
which thin out distally and give the ledge a blunt free edge.
The ledge has a thickness, proximally, of nearly 0.08 oral
diameter. The bowl, below the collar, swells out to its
maximum diameter of 1.3 1 oral diameters at 0.73 oral diam-
eter below the rim. The sides of the bowl are full and some-
what lumpy locally in contour. The aboral end is evenly
rounded to blunt (no°) and without any special differentia-
tion. The end is closed.

The wall is relatively thick, being 0.07 oral diameter in
thickness across the bowl. In the collar region it thins down
rapidly to the sharp-edged oral rim. The wall has a distinct,
thin inner lamina which coats the interior of the collar and
bowl. There are thin-walled, large, radial secondary poly-
gons in a single layer in the bowl; in the ledge these are in
two rows, and just below the oral margin there are minute
ones in several rows. There are about 33 hexagons across the
bowl before the ledge and 24 from ledge to aboral end.
Those immediately below the rim are minute and become
larger at the rim. Below the ledge they are subuniformly still
larger. The wall is soft and flaccid and is easily flattened by
pressure; the texture is not at all like that of Cyttarocylis.

Length, 85 to 90(1.

Loricae of Codonopsis ollula have been obtained only in a
limited area of the Pacific under rather uniform external
conditions. There seems to be little difference between
them, save, perhaps, in the shape of the aboral ends, and this
difference may be due to cover-slip pressure when the speci-
mens were examined in formalin-sea water under rather
heavy, long slips.

Codonopsis ollula is unique. It bears some superficial
likeness to Cyttarocylis longa, but the wall structure is alto-
gether different, and the ledge of ollula is sufficient to dis-
tinguish it from any Cyttarocylis.

Kofoid and Campbell (1929) followed Brandt (1907) in
the assignment of this species to Cyttarocylis. but later ( 1939)
these authors reassigned it to a new genus, Codonopsis, in
which it is here retained.

Recorded from five stations in the Pacific, as follows: one
(47) in the Galapagos region, one (48) in the region of

TAXONOMY AND DISTRIBUTION

*7

South Pacific island fields, one (54) in the South Pacific-
middle latitudes, one (99) in the Pacific equatorial region,
and one ( 100) in the North Pacific trade region. Codonopsis
(Alula has a limited distribution in the central Pacific.

There are 5 net samples, of which 2 were taken at the
surface, 1 at 50 meters, and 2 at 100 meters. Maximum
frequency, 4 per cent at stations 48, 54; elsewhere, minimum
only; average, 2.5 per cent.

Temperature, i8?74~27?84 (24:35); salinity, 34.71-36.44
(35.50): density, 22.31-25.37 (23. 89); pH, 8.16-8.23 (8.21).

CYTTAROCYLIDAE Kofoid and Campbell

Cyttarocylidae Kofoid and Campbell, 1929, p. 108.

Only one genus is included in this family, and this one is
allied to the higher members of the Codonellidae. It is
exclusively eupelagic, mostly in warmer seas.

CYTTAROCYLIS Fol emended
Cyttarocylis, Kofoid and Campbell, 1929, p. 109.

Cyttarocylis is an isolated genus with some alliance to
Codonella, with which it agrees in general, although more
complex and finished in form and wall structure.

Cyttarocylis is widely distributed in warm seas as well as
in more temperate waters. It does not enter the Arctic
Ocean, although Cyttarocylis magna occurs in Mutsu Bay to
the north, and as far south as below Australia. The majority
of the species are tropical but not limited to either of the
oceans investigated in this report.

Nine species are described.

Cyttarocylis acutiformis Kofoid and Campbell

(Figures 35, 36, 37, 45, 48)

Cyttarocylis acutiformis Kofoid and Campbell, 1929, p. ill, fig.
221.

The elongated, conical lorica, with flaring, distinct collar
and pointed aboral end, has a length of 1.78 to 2.35 (2.07)
oral diameters. The oral margin is minutely denticulate
with about 70 triangular, sharply pointed denticles, ragged
in a few instances, or with few larger, outward-directed
teeth. The collar is a basal segment of an inverted truncated
cone (40 to 50 ), with a length of 0.12 to 0.15 total length.
The sides of the collar are usually flattened or, in very per-
fect individuals, flat sigmoid. The nuchal shelf reduces the
aperture between collar and bowl to 0.77 oral diameter. It
is triangular and horizontal, with barely concave sides. The
long, conical bowl is slightly convex (about 30 ); it is 25
near the throat and for 0.67 of the length, and 35 ° in the
aboral region. The latter region is sharply pointed to
minutely truncated and always ends as an open canal which
in a few individuals may be relatively wide open.

The wall has about 56 to 60 polygons across one face and
75 to 90 from rim to tip. These polygons are mostly minute,
and prevailingly pentagonal to hexagonal, with some tri-
angular ones. Large and small reticulations are inter-
mingled, but, in general, they are larger below the throat

and in the upper three-tenths ol the bowl. There are one or
two rows of large polygons in the collar and several rows of
smaller ones, especially immediately below the oral rim.
The wall is thickest in the upper part of the bowl, being 0.04
oral diameter at the most. It is only barely thinner in the
lower bowl. Only a single layer of rectangular prisms occurs
except in the nuchal shelf, where there are two layers of pen-
tagons. The cavity of the lorica conforms to the outer con-
tour closely save as the nuchal shelf constricts the opening.
The narrow aboral canal is usually a tube.

There are 20 macronuclei, and zooxanthellae are present.
The membranelles are large blades.

Length, 213 to 326 (228.6) p..

The five loricae figured are extremes picked out from a
large number of specimens. The oral margin, flare of collar,
shape of bowl, and shape of aboral end are the principal
characters subject to variation. These loricae are longer and
less stout than those originally described by Kofoid and
Campbell, which are 200 to 257P in length, 1.54 to 1.96 oral
diameters. Possibly these came from cooler waters. The
fine meshwork is a most uniform character.

Cyttarocylis acutiformis is most like C. magna, but is
generally smaller, with a more slender bowl, a more clearly
flaring collar, and finer reticulations. It resembles C. cassis,
but is greatly elongated and less convex, and lacks a terminal
peg; its surface is more finely reticulated. The aboral end
is not truncated like that of C. conica, and the bowl is not
so full. It lacks the aboral contraction of both C. mucronata
and C. obtusa.

Recorded from thirty-one stations, twelve in the Atlantic
and nineteen in the Pacific, as follows: two (2, 16) in the
Gulf Stream, two (17, 20) in the Sargasso Sea, seven (22, 23,
24, 25, 27, 28, 30) in the Atlantic equatorial region, one (34)
in the Caribbean Sea, six (45, 46, 47, 75, 78, 80) in the Gala-
pagos region, two (84, 85) in the region of South Pacific
island fields, one (64) in the South Pacific middle latitudes,
three (113, 144, 145) in the North Pacific middle latitudes,
five (131, 137, 146, 148, 149) in the California region, and
two ( 150, 151 ) in the North Pacific trade region.

There are 7 pump and 28 net samples, of which 2 were
taken at the surface, 18 at 50 meters, and 15 at 100 meters.
Maximum frequency 43 per cent at station 23 at 100 meters;
other records above minimum (2 to 42 per cent) from sta-
tions 2, 16, 17, 22, 23, 25, 27, 28, 30, 34, 45, 75, 80, 131, 137,
145, 146, 148, 149, 150, 151; averages, 1.5 and 1.0 loricae in
Atlantic and Pacific pump samples, and 11.6 and 3.6 per cent
in Atlantic and Pacific net samples, respectively.

Temperature: Atlantic, pump samples i4?6o-2i?49( i8?04),
net samples i4?6o-36?02 (22^07); Pacific, i7?46-26?o6
(2i?52) and i2?i2-27?89 (2t?48), respectively. Salinity:
Atlantic, pump samples 35.70-36.02 (35.86), net samples
35.61-36.73 (36.21); Pacific, 34.42-36.17 (35.34) and 34.32-
36.42 (35.14), respectively. Density: Atlantic, pump samples
25.15-26.62 (25.88), net samples 23.79-26.62 (25.24); Pacific,
23.42-25.11 (24.43) anc ^ 22-48—25.55 (24.75), respectively.
pH: Atlantic, pump samples 7.93-8.22 (8.07), net samples
7.93-8.30 (8.18); Pacific, 8.14-8.33 (8.22) and 8.10-8.38
(8.34), respectively.

i8

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Cyttarocylis brandti Kofoid and Campbell

(Figure 39)

Cyttarocylis brandti Kofoid and Campbell, 1929, pp. 111-112,

fig. 215.
Cyttarocylis plagwstoma, Hofker, 1931, pp. 37 I_ 37 2 . "g s - 5 I_ 53-

The aborally blunt lorica is acorn-shaped and 0.87 to 0.98
oral diameter in length. The oral margin is finely denticu-
late. The collar is a short basal segment of an inverted
truncated cone (90° to ioo°) with a length of 0.1 oral
diameter. Its sides are slightly convex. The nuchal shelf
is a thin, upturned (50 ) ridge at the throat. The bowl is
baggy and convex conical (about 35 ° at the upper end and
130 near the apex). The aboral end is blunt, barely pointed,
and usually undifferentiated.

The wall has about 75 small polygons below the collar
and about 40 from throat to tip. The polygons are prevail-
ingly hexagons and a few large ones are scattered near the
upper end of the bowl. The bowl is thickest near the shelf,
being less than 0.0 1 oral diameter, and has only a single
layer of more or less subuniform secondary prisms. The
lumen follows the outer contour save as the nuchal shelf
alters it.

Length, 90 to i2op_.

The lorica figured (fig. 39) differs from the usual type in
the shape of the bowl, and also in having a tiny nipple. Its
nuchal shelf is almost horizontal.

Cyttarocylis brandti resembles C. longa in being generally
more pointed, and in having a less baggy bowl. It is usually
longer than C. eucecryphalus, and that species has a flat
aboral end. The bowl of C. brandti is less conical than that
of C. cassis, and proportionately stouter. It is most likely to
be confused with longa.

Hofker (1931) figures 2 loricae called Cyttarocylis plagi-
ostoma, but these appear to belong to the present species.
The general proportions, shape of collar, and aboral end are
the same, as is also the wall pattern. Hofker finds 18 mem-
brandies with no intercalary platelets, and records 80 macro-
nuclei. He suggests, probably correctly, that most of these
are zooxanthellae. Careful cytological investigation will prob-
ably show that most members of this genus are photosynthetic.
Recorded from thirty-seven stations, nine in the Atlantic
and twenty-eight in the Pacific, as follows: four (18, 19, 20,
21) in the Sargasso Sea, five (22, 23, 24, 25, 27) in the
Atlantic equatorial region, four (41, 42, 45, 75) in the
Galapagos region, four (54, 55- 6 5* 6 7) in the Soutn Pacinc
middle latitudes, three (84, 95, 96) in the region of South
Pacific island fields, four (101, 140, 150, 151) in the North
Pacific trade region, three (115, 141, 145) in the North
Pacific middle latitudes, eight (131, 132, 135, 136, 146, 147,

148, 149) in the California region, and two (153, 154) in
the Pacific equatorial region.

There are 9 pump and 36 net samples, of which 11 were
taken at 50 meters and 34 at 100 meters. Maximum fre-
quency, 4s per cent at station 153; other records above mini-
mum (2 to 37 per cent) from stations 18, 19, 20, 22, 23, 27,
41, 45, 54, 55, 65, 67, 75, 115, 131, 136, 14s. 146. 147. J 4 8 >

149, 150, 151, 154; averages, 1.0 lorica in both Atlantic and

Pacific pump samples, and 6.0 and 12.5 per cent in Atlantic
and Pacific net samples, respectively.

Temperature: Atlantic, pump sample 2i?49, net samples
i4?6o-36?02 (22^29); Pacific, i4?42-28?4i (20^04) and
i2?i2-28?74 (i8?76), respectively. Salinity: Atlantic, pump
sample 36.02, net samples 35.61-37.05 (36.26); Pacific, 33.40-
35.63 (34.80) and 33.36-36.42 (34.87), respectively. Density:
Atlantic, pump sample 25.15, net samples 23.98-26.64 (25.29);
Pacific, 22.75-24.88 (24.05) and 22.98-26.11 (24.91), respec-
tively. pH: Atlantic, pump sample 8.22, net samples 7.93-
8.25 (8.15); Pacific, 8.19-8.34 (8.29) and 7.91-8.39 (8.22),
respectively.

Cyttarocylis cassis (Haeckel) Fol
Cyttarocylis cassis, Kofoid and Campbell, 1929, p. 112, fig. 219;

Alzamora, 1929, p. 11, fig. 24; Hofker, 1932, p. 370, figs. 49.

50.
Tintinniis sp., Lindeman, 1924, p. 889, fig. 10.

The conical lorica, with widely flaring collar and pointed
aboral end, has a length of 1.43 oral diameters. The oral
margin is slightly ragged and there are minute teeth around
its edge. The short collar is a basal segment of an inverted
truncated cone (50 ), and its length is about 0.08 total
length; it is well developed and flares widely. Its sides are
mildly convex. The lower edge is marked by an optically
dark line. The nuchal shelf is angular (90 ) and horizontal,
and barely narrows the aperture between collar and bowl.
The full conical bowl changes from about 14 in the an-
terior three-fifths to 52 in the aboral section. There is a
tiny but distinctive nipple-like aboral peg at its free tip. The
bowl as a whole is decidedly convex.

The wall is coarsely reticulated, with about 30 polygons
across the throat and about 36 from oral rim to tip. The
polygons are prevailingly rounded, and some are pentagons.
They are of various sizes, small ones being scattered among
the larger. Those of the upper half, including the collar, are
larger than those of the lower part of the bowl near the
aboral end, where they are much smaller and more nearly
circular. The wall has a subuniform thickness of 0.03 oral
diameter, except at the nuchal shelf, where it is 0.05. Single
layers of polygons occur. The cavity of the lorica is altered
by the nuchal shelf. The aboral peg is solid.
There are 18 to 20 macronuclei.
Length, 170 to 215(1.

The Carnegie specimens hardly differ from the usual form
found in the Mediterranean.

Cyttarocylis cassis differs from C. mucronata and C. obtusa
in having a convex-conical bowl instead of one narrowed
down to a pedicel-like aboral region. It lacks the aboral
flattening of C. conica. It is less elongated and more convex
conical than C. acutiformis; it is also stouter and shorter, and
has a terminal peg. Cyttarocylis magna is longer and more
slender and has a less distinct flaring collar. Cyttarocylis
cassis is long familiar, and recent records conform to type.
Recorded from twenty-two stations, twelve in the Atlantic
and ten in the Pacific, as follows: one (2) in the Gulf
Stream, two (17, 18) in the Sargasso Sea, seven (22, 23, 24,
25, 27, 28, 29) in the Atlantic equatorial region, two (31, 32)

TAXONOMY AND DISTRIBUTION

19

in the Caribbean Sea, three (55, 65, 67) in the South Pacific
middle latitudes, three (68, 75, 80) in the Galapagos region,
two (131, 140) in the California region, and two (140, 151)
in the Pacific trade region.

There are 29 net samples, of which 1 was taken at the
surface, 6 at 50 meters, and 22 at 100 meters. Maximum
frequency, 7 per cent at station 27; other records above
minimum ( 2 to 6 per cent) from stations 2, 17, 22, 23, 24,
28, 32, 75, 80, 131; averages, 3.0 and 1.6 per cent in the
Atlantic and Pacific, respectively.

Temperature: Atlantic, i4?6o-26?97 (20^97); Pacific,
i2 L :i2-26?o6 (i8?7o). Salinity: Atlantic, 35.11-36.81 (36.14);
Pacific, 34.30-35.95 (34-7 2 )- Density: Atlantic, 23.98-26.62
(25.40); Pacific, 22.80-25.52 (24.83). pH: Atlantic, 7.93-
8.26 (S.i(>); Pacific, 8.09-8.39 (8.17).

Cyttarocylis conica Brandt
(Figure 40)
Cyttarocylis conica, Kofoid and Campbell, 1929, pp. 112-113,

fig. 220.
The truncated-conical lorica has a length of 1.47 oral diam-
eters. The oral margin is irregular and ragged. The collar
forms a short segment of an inverted truncated cone (48 )
with even, regular sides, and its length is 0.12 total length.
The nuchal shelf is developed inside at its lower edge. The
shelf is feebly developed, angled, and horizontal. The gen-
erally conical bowl is formed below the shelf and there is
almost no nuchal constriction. The bowl has a length of
0.88 total length. It changes from about 22 suborally to
6o° in the aboral region and has well filled sides. The aboral
end is squarely truncated, and about 0.18 oral diameter in
diameter. At the middle of the aboral end there is a minute
conical aboral peg, which is closed.

The wall is regularly and evenly reticulated, with 50 or
more polygons around the suboral region and 35 to 42
around the meridian. The polygons are prevailingly pen-
tagonal to hexagonal, and subequal; each has an enclosing
mesh with thickened, heavy beams. The wall is about 0.04
oral diameter in thickness just below the throat, at the
thickest level, and elsewhere is thinner. The large secondary
prisms occur always in a single layer.
Length, 15711.

The Carnegie loricae are somewhat shorter than those
recorded elsewhere, the latter reaching to 21 on. In propor-
tions and general shape the agreement with others is, how-
ever, close, and the smaller specimens may have come from
warmer water than others from the Atlantic.

This species is unlike any of the others in the shape of the
aboral end, there being aboral flattening. For this reason
there can be little opportunity to confuse it with others. In
Cyttarocylis cassis the aboral end is pointed and the whole
bowl is a broad cone.

Recorded from two stations in the Atlantic, 19 and 20,
both in the Sargasso Sea.

There are 2 net samples. Frequency, minimum.
Temperature, 22°42-22?56 (22^49); salinity (one record),
37.05; density (one record), 25.67; pH, 8.18-8.25 (8.21).

Cyttarocylis eucecryphalus (Haeckxl) Koloid
(Figure 42)

Cyttarocylis eucecryphaltts, Kofoid and Campbell, [929, p. 113,

fig. 211.
The stout, acorn-shaped lorica, with llaring collar and flat
aboral end, has a length of 1.0 oral diameter, The oral mar-
gin is regularly beset with numerous subequal, equidistant,
triangular, pointed teeth. The collar is a basal segment of
an inverted truncated cone (57°), with a length of 0.14 total
length. Its sides are slightly convex. The nuchal shelf is a
concave, virtually horizontal triangle (4} ), and its width
is such that the diameter of the aperture between collar and
bowl is reduced to 0.72 oral diameter. The bowl is baggy,
increasing from about 12 in the upper 0.5 to 58 in the
lower section; nearer the aboral end it increases to 100°.
The aboral end is flattened, although not squarely truncated,
and its diameter is 0.32 oral diameter. There is no point.
The wall is coarsely reticulated with about 23 polygons
across the throat and 17 from rim to tip. There is but a
single row around the collar. The polygons are largest in
the upper half of the bowl and become progressively smaller
in the lower part. They are prevailingly pentagonal, al-
though some are rounded triangles and some are hexagons
with rounded edges. The wall is subuniformly 0.03 oral
diameter in thickness, and the rectangular prisms occur in a
single layer in the gray, semitransparent wall. The lumen
is reduced by the internal shelf but otherwise follows the
outer contour.

Length, 1 15 to 140^1.

A few loricae tend toward a thimble-like bowl like the
one figured (fig. 42).

Cyttarocylis eucecryphalus is closely related to C. brandtt,
but its aboral end is flattened instead of blunted. It is like
C. longa, but its aboral end, again, is different. It is more
slender than C. ricta and has coarser meshwork. Cyttarocylis
plagiostoma is sharply pointed and has yet coarser meshwork.
Recorded from forty-eight stations, sixteen in the Atlantic
and thirty-two in the Pacific, as follows: two (2, id) 111 the
Gulf Stream, one (4) in the Atlantic drift, four (17, 18, 19,
20) in the Sargasso Sea, eight (22, 23, 24, 25, 26, 27, 28, 29)
in the Atlantic equatorial region, one (33) in the Caribbean
Sea, eleven (41, 42, 45, 46, 47, 69, 75, 77, 78, 79, 80) in the
Galapagos region, five (48, 49, 81, 82, 95) in the region ol
South Pacific island fields, three (54, 66, 67) in the South
Pacific middle latitudes, two (113, 144) in the North Pacific
middle latitudes, seven (131, 135, 136, 137, 146, 147, 149) in
the California region, and four (109, 139, 150, 151) in the
North Pacific trade region.

There are 20 pump and 53 net samples, of which 7 were
taken at the surface, 32 at 50 meters, and 34 at 100 meters.
Maximum frequency, 86 per cent at station 17 at 100 meters;
other records above minimum (2 to 74 per cent) from sta-
tions 2, 16, 18, 19, 20, 22, 23, 24, 25, 26, 27, 28, 33, 41, 45, 46,

47, 48, 75- 77- 7 8 - 8 °. 8l - 95- i°9, 131. 136, J37- '39. 146, M7.
149, 150; averages, 1.5 and 18.0 per cent in Atlantic pump
and net samples, and 1.0 and 4.8 per cent in Pacific pump
and net samples, respectively.

20

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Temperature: Atlantic. pumpsamples20?99-26?98 (23^28),
net samples i2?i2-28? 74 (2i?85); Pacific, i 4 ?77-26?7o
(2i?58) and i3?37-26?79 (2i?29), respectively. Salinity:
Atlantic, pump samples 36.04-37.00 (36.50), net samples
33-36-3 6 44 (35-22); Pacific, 34.59-36.04 (35.36) and 35.61-
37.15 (36.38), respectively. Density: Atlantic, pump samples
24.25-25.58 (24.95), net samples 22.43-26.17 (25.47); Pacific,
23.42-25.91 (24.02) and 23.70-27.01 (25.43), respectively.
pH: Atlantic, pump samples 8.14-8.27 (8.22), net samples
7.91-8.39 (8.20); Pacific, 8.10-8.39 (8.26) and 7.93" 8 - 2 7
(8.23), respectively.

Cyttarocylis longa Kofoid and Campbell
(Figures 41, 43)
Cyttarocylis longa Kofoid and Campbell, 1929, p. 113, fig. 217.
The stout, saccular lorica, with rounded to barely pointed
aboral end, has a length of 0.76 to 1.08 oral diameters. The
oral margin is minutely denticulate and irregular. The
collar is a basal segment of an inverted truncated cone (65
to 75 ) with a length of 0.12 to 0.15 oral diameter. The
nuchal shelf is horizontal, narrow, and concave on the
superior face. The bowl is sac-shaped, with its greatest
diameter at the throat, 30 suborally and 115 to 130°
aborally. The aboral end is evenly rounded to blunt; rarer
instances show a trace of aboral pointing.

The wall is finely reticulated with triangular, pentagonal,
or hexagonal polygons. There are 40 to 60 polygons around
the throat and 32 to 45 from oral rim to tip. In general, the
polygons are graded in size from larger to smaller from the
rim downward. There is a double row of large ones with
interspersed tiny ones in the collar. The wall has a subuni-
form thickness of 0.03 oral diameter in the bowl. The wall
of the collar thins out from the internal ledge to the rim.
There is a single layer of prisms in the wall, and interpris-
matic substance is pronounced between the prisms.

Length, 103 to io6u.

There is considerable variation in the shape of the aboral
end, which may be evenly rounded, blunt, or rarely pointed,
but there is never a nipple. The oral rim is rarely ragged.
The Carnegie loricae are 0.76 to 1.08 oral diameters, as
against 0.76 to 0.96 recorded by Kofoid and Campbell
(1929).

Cyttarocylis longa differs from C. brandti in being longer,
more saccular in bowl, and less evidently pointed, and in
having much finer reticulations. It differs from C. eitce-
cryphalus in the shape of the aboral end, that of the latter
species being flat. The wall of the two species is also dif-
ferent.

Cyttarocylis plagiostoma has a fuller, sharply pointed
bowl and a coarse surface, is longer and less stout, and has
better separation of collar and bowl than is found in C. ricta.
The latter is most likely to be confused with brandti.

Recorded from thirty-two stations, nine in the Atlantic
and twenty-three in the Pacific, as follows: one (2) in the
Gulf Stream, seven (22, 23, 24, 25, 26, 27, 30) in the Atlantic
equatorial region, one (34) in the Caribbean Sea, eight (46,
47, 68, 70, 72, 77, 78, 80) in the Galapagos region, three (48,

159, 160) in the region of South Pacific island fields, three
(54, 63, 67) in the South Pacific middle latitudes, two (109,
151) in the North Pacific trade region, four (131, 136, 147,
148) in the California region, one ( 145) in the North Pacific
middle latitudes, and two ( 153, 156) in the Pacific equatorial
region.

There are 10 pump and 30 net samples, of which 5 were
taken at the surface, 11 at 50 meters, and 24 at 100 meters.
Maximum frequency, 29 per cent at station 46; other records
above minimum ( 2 to 27 per cent) from stations 2, 22, 27,
47, 48, 54, 67, 70, 78, 109, 131, 136, 145, 147, 151, 153, 156,
159, 160; averages, 1.5 and 6.0 per cent in Atlantic pump
and net samples, and 1.0 and 7.9 per cent in Pacific pump
and net samples, respectively.

Temperature: Atlantic, pump samples i8?40-26?04(i8?72),
net samples i4?6o-36?o2 (22?33); Pacific, i2?73~24?38
(20^52) and i2?i2-28?52 (2^24), respectively. Salinity:
Atlantic, pump samples 36.08-36.25 (36.16), net samples
35.61-36.58 (36.09); Pacific, 34.58-36.04 (35-50) and 33.36-
36.04 (35.30), respectively. Density: Atlantic, pump samples
23.98-26.01 (24.99), net samples 23.26-26.62 (25.22); Pacific,
24.33-26.28 (24.99) and 22.76-26.28 (24.50), respectively.
pH: Atlantic, pump samples 8.11-8.30 (8.20), net samples
7.93-8.30 (8.13); Pacific, 7.68-8.19 (8.09) and 7.68-8.44
(8.21), respectively.

Cyttarocylis magna Brandt

(Figures 44, 46, 47)

Cyttarocylis magna, Kofoid and Campbell. 1929, p. 114, fig.
222; Hada, 19326, p. 564, fig. 16.

The tall, conical lorica, with low, ill defined collar and
short, pedicellated aboral end, has a length of 2.41 to 3.18
(2.77) oral diameters. The oral margin is finely to roughly
ragged. The short collar is a basal segment of an inverted
truncated cone which rarely flares as much as 40 , and has
a length of 0.10 to 0.15 oral diameter. The collar is poorly
or scarcely at all delimited from the long bowl. The nuchal
shelf is triangular in section, horizontal, and feebly devel-
oped. The sides of the collar are flat to convex, usually the
former. The full conical bowl changes from about 25 ° below
the collar to 45 near the apex, and there is a small, conical
aboral horn (25 ) with a length of only 0.1 oral diameter.

The wall is coarsely reticulated with subuniform hexagonal
polygons. There are about 25 of these across the face at the
throat and 45 to 50 from the throat to the apex. Those of
the collar do not differ from those of the throat, and they
are only slightly smaller aborally. The wall is about 0.0 1
oral diameter in thickness. There is but a single layer of
polygons, even in the nuchal shelf. The lumen follows the
outer contour neatly save as the low internal shelf breaks the
collar and bowl into two divisions. The aboral horn is
hollow and the tip is usually open.

Length, 267 to 400U.

There is considerable variation in these large loricae. In
actual length, Kofoid and Campbell (1929) record 250 to
320ft and Hada (19326) 300(1; the Carnegie specimens are
267 to 400JI, and in proportions also they are more variable.

TAXONOMY AND DISTRIBUTION

21

In form, these conform to the general shape of others; the
collar is sometimes more definitely separated from the bowl,
and the horn is lacking in a few.

Cyttarocylis magna is most nearly like C. acutiformis, but
is larger, being up to 400U as against a maximum of 326^1;
its proportions are also different and there is less distinction
between collar and bowl. The ragged margin and aboral
horn are added differences. It differs in these characters
from C. cassis, and also in being longer and less stout, and
h.i\ ing a less flaring collar. It is unlike the other species and
can seldom be confused with them.

Recorded from twelve stations, four in the Atlantic and
eight in the Pacific, as follows: one ( t6) in the Gulf Stream,
two ( 17, 18) in the Sargasso Sea, one (27) in the Atlantic
equatorial region, two (35, 35-36) in the Pacific equatorial
region, three (45, 70, 7s) in the Galapagos region, one (67)
in the South Pacific middle latitudes, and two ( 109, 151) in
the North Pacific trade region.

There are 14 net samples, of which 1 was taken at the
surface, 5 at 50 meters, and 8 at 100 meters. Maximum fre-
quency, 32 per cent at station 35; other records above mini-
mum (2 to 25 per cent) from stations 35-36, 45, 67, 70, 7=5,
151; average in the Pacific, 9.2 per cent; in the Atlantic there
were only minimum occurrences.

Temperature: Atlantic, i8?o8-23?64 (2i?2o); Pacific,
i4?33~2i?69 (17:24). Salinity: Atlantic, 36.03-36.82 (36.53 ):
Pacific, 34.42-35.47 (35.02). Density: Atlantic, 24.84-26.07
(26.00); Pacific. 24.48-26.28 (25.36). pH: Atlantic, 8.09-
S.27 (8.20); Pacific, 7.68-8.18 (7.98).

Cyttarocylis mucronata Kofoid and Campbell

(Figure 38)

Cyttarocylis mucronata Kofoid and Campbell, 1929, p. 114, fig.
216.

The elongated, acorn-shaped lorica, with contracted aboral
cone and aboral peg, has a length of 1.56 oral diameters.
The oral margin is coarsely irregular and rather ragged.
The short collar forms a segment of an inverted truncated
cone (50 ), and has evenly convex, full sides. Its length is
only 0.1 1 total length. The nuchal shelf is developed inside
at its lower edge. The shelf is thin, angled, upright, and not
well developed. The generally conical bowl is formed below
the clear-cut collar. The bowl is subdivided into a long sub-
oral inverted truncated conical section (20 ), with a length
of 1.0 oral diameter, and an aboral section which is a shorter,
wider inverted cone (62 ) with a length of 0.4 oral diam-
eter. The sides are barely convex. The aboral portion is a
pedicel-like inverted cone (58°) with a length of 0.2 oral
diameter and with convex sides. At its posterior end is a
tiny aboral peg.

The wall has about 60 to 65 prevailingly hexagonal, rather
fine polygons around the throat, and 45 or more from oral to
aboral end. The polygons are larger and subuniform near
the rim, and become progressively smaller below. The wall
has a subuniform thickness of less than 0.02 oral diameter.
There is a single layer of secondary prisms. The aboral peg
is solid but otherwise the cavity follows the outer contour.

Length, 152ft.

The lorica figured (fig. 38) differs from that figured by
Kofoid and Campbell (1929) in having a coarsely ragged
oral rim and in being a little longer.

Cyttarocylis mucronata differs from C. obtusa in its finer
surface reticulations and in the presence of an aboral peg.
This terminal point is distinct and invariable. The aboral
end of obtusa is blunt and squarely cut and lacks a peg. The
aboral cone of both species lacks a parallel in the genus. The
aboral truncation of C. mucronata is like that of C. conica,
but that species is longer and more slender as well as dif-
ferent in other ways.

Recorded from four stations in the Atlantic, as follows:
two (19, 20) in the Sargasso Sea, and two (22, 25) in the
Atlantic equatorial region.

There are 4 net samples, all taken at 100 meters. Maxi-
mum frequency, 3 per cent at station 19: other records at
the minimum; average, 1.6 per cent.

Temperature, i4?6o-22?56 (i9?27); salinity, 35.70-37.05
(36.49); density, 25.38-26.67 (25.69); pH, 7.93-8.25 (8.12).

Cyttarocylis obtusa Kofoid and Campbell
Cyttarocylis obtusa Kofoid and Campbell, 1929, p. 115, fig. 218.

The elongate, rather slender, acorn-shaped lorica. with
contracted aboral cone and flattened end without peg, has a
length of 1.06 to 1.36 oral diameters. The oral margin is
minutely denticulate. The short collar forms a segment of
an inverted truncated cone (90 ), and has a length of 0.1
total length. Its sides are flat except just below the margin,
where they turn upward. The nuchal shelf is inconspicuous,
upright, and thin-edged. The bowl is quite convex conical
(30 ) in the suboral seven-tenths, and contracted (8o°)
below. Its sides are slightly sinuous. The aboral subdivision
is a flat-ended cone (36 ) and its aboral end has a diameter
of 0.14 oral diameter. Its sides are even. There is no trace
of an aboral peg.

The wall has about 36 to 40 polygons around the neck
and 37 to 42 from oral edge to tip. The polygons are pre-
vailingly hexagonal, and larger in size in the suboral three-
tenths than below; in the posterior section they are only one-
third to one-half the size of those superior, and they tend to
be round. The wall is subuniformly 0.01 oral diameter in
thickness at the nuchal shelf, and there is but a single layer
of prisms. The cavity neatly conforms to the outer contour.

Length, 135 to 157(1.

Significant variations in length are probably correlated
with temperature adjustment.

Cyttarocylis obtusa differs from C. mucronata in the dis-
tinct lack of an aboral peg, and in the generally smaller sur-
face polygons, more regular oral rim, and slimmer loricae.
The aboral end of C. conica is also truncated, but that species
has no aboral cone and the proportions are unlike, being 1.41
to 1.47 in conica.

Recorded from three stations in the Atlantic, as follows:
two (18, 19) in the Sargasso Sea, and one (22) in the
Atlantic equatorial region.

There are 3 net samples, of which 1 was taken at 50

22

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

meters and 2 at 100 meters. Maximum frequency, 5 per cent

at station 22; other records minimum; average, 3 per cent.

Temperature, i7?5o-25?3i (2i?04>; salinity, 36.81-37.15

( 56.98); density, 24.89-26.07 (25.48); pH, 8.21-8.27 (8.24).

CODONELLOPSIDAE Kofoid and Campbell
Codonellopsidae Kofoid and Campbell, 1929, p. 67.
This family includes three genera, Stenosemella, Codonel-
lopsis, and Laacknianniella , of which the first two occur in
the material of this expedition. Laac\manniella is exclu-
sively antarctic, Codonellopsis is eupelagic in warm seas, and
Stenosemella is almost always coastal.

STENOSEMELLA )6rgensen
Stenosemella, Kofoid and Campbell, 1929, pp. 67-68.

Stenosemella is apparently the stem of the Codonellopsidae,
and its wall structure allies it to Tintinnopsis, to the general
form of which it adds a collar. Closely related is the new
genus Wangiella, with the type, Wangiella dicollaria, from
the Bay of Amoy (D. Nie, 1934). in which the collar is
Dictyocysta-Mke. Also allied is Luminella, a new genus
founded by Kofoid and Campbell (1939, p. 284), which has
as its principal character the demilunar openings in the
collar.

Stenosemella is rare in the open ocean except as drifting
empty loricae may be carried from the coast; in harbors and
close to shore it is frequently the most common tintinnid.
In San Francisco harbor it is often found exclusively, and
off La Jolla it is often the most abundant. Most records of
the genus are from northern European waters.

One species is described here.

Stenosemella nivalis ( Meunier) emended Kofoid
and Campbell
Stenosemella nivalis, Kofoid and Campbell, 1929, pp. 69-70, fig.
136; Campbell, 1931, pp. 347"34 8 ; Hada - r 93^>. P- 561, fig-
11; Marshall, 1934, p. 638.
Stenosemella nucula, Hofker, 1931, pp. 362-364, figs. 40-42.

The small, olive-shaped lorica, with thick, irregular wall
and narrow, hyaline collar lacking windows, has a length of
2.2 oral diameters. The oral margin is recurved outward,
and the glassy collar is a concave ring with a length of only
0.06 oral diameter; it lacks the spiral turns and demilunes
characteristic of some species, and the tall windows of
Wangiella dicollaria. The plump, dense, potlike bowl is in-
flated from the lower edge of the collar and reaches its
greatest diameter, which is equal to the total length, at 0.61
oral diameter below the oral rim, forming a rather strong,
rounded shoulder. Below this level it contracts in regular
contour to the bluntly pointed or narrowly rounded end.

The wall of the collar is clear, but that of the bowl is
coarse, thick, and heavy, up to 0.2 oral diameter laterally.
No structure may be detected in the collar, but the bowl has
two or three layers of large, irregular tertiary polygons with
thickened margins and clear interiors. Foreign bodies adhere
to the outer surface.

The animal has been carefully studied. There are 2
macronuclei, and 24 large, squarish membranelles of elabo-
rate structure (Campbell, 1931).
Length 40 to 46^.

Considerable variation is apparent, especially in the rela-
tive narrowness or width of the bowl, the level at which the
greatest diameter is reached, the shape of the aboral end, and
the character of the collar. Cytological details also differ.
Stenosemella nivalis is closest to S. ventricosa in form, but
nivalis is always shorter, and rotund and sharper aborally.
The greatest diameter is reached at a level closer to the
collar in ventricosa. Stenosemella nivalis has a different
collar from Wangiella dicollaria from the Chinese coast; it
is smaller than S. steini, and lacks the trough below the
collar which distinguishes that species. Stenosemella avellana
is of about the same size, but its greatest diameter is nearer
the middle and there are long, sloping shoulders below the
collar. Stenosemella oliva has a narrower bowl and higher
collar, and is differently proportioned.

Hofker ( 1 93 1 ) suggests that the western species is nivalis,
ami that the loricae from off Holland are nucula. This
proposition is largely based on inadequate cytological mate-
rial. In form, loricae from Holland as examined by us seem
similar in every way to those from off California.

Recorded from one station (130) in the California region,
in a net sample taken at 50 meters. Frequency, 5 per cent.
Temperature, 12^91; salinity, 33.40; density, 25.19; pH,
8.26.

CODONELLOPSIS JSrgensen
Codonellopsis, Kofoid and Campbell, T929, p. 73.
Stenosemella may be considered the stem of Codonellopsis,
and Luminella that of Dictyocysta. Both these primitive
genera are probably derived from Tintinnopsis. This hy-
pothesis leaves the more highly differentiated genera less
isolated.

Codonellopsis is widely distributed in the warmest oceans,
although a few species enter the cold North Pacific, for
example, C. frigida, C. limosa, and C. orientalis northward
of Japan, and the latter and aleutiensis southward of the
Bering Sea. A few, such as C. contracta, also occur in the
tropics in deep water. None of the species enters the Ant-
arctic, but the allied genus Laac\manniella is peculiar to
that region. Laae\manniella has two, or possibly more,
species peculiar to the south, where they occur under the
ice; this genus does not occur in the Humboldt Current,
although the water of that current arises from the Antarctic
Current. It has not been reported, either, from the Cape
Horn Current. Laacl{manniella is remote from Coxliella
and related genera, and belongs properly to the Codonellop-
sidae. Its open aboral end is secondary. Most species of
Codonellopsis are circumtropical.

Nineteen species are described here, of which one is new.

Codonellopsis aleutiensis, new species
(Plate 1, figure 9}
The lorica has a long collar and a round bowl; its length
is 2.66 oral diameters. The oral margin is entire, rounded

TAXONOMY AND DISTRIBUTE )\

23

over, and comparatively thin. The long collar is subdivided
into a short anterior widely flaring funnel, and a longer sub-
cylindrical posterior section. The suboral funnel is an in-
verted truncate cone (28 ). Its sides are full, although they
do not bulge, and the length is 0.17 total length, or about
0.52 oral diameter. The funnel includes the upper 4 turns
of the spiral lamina which forms the collar. The remaining
part of the collar is a truncated cone (about 7 ). Its diam-
eter at the oral end where it joins the suboral funnel is 0.78
oral diameter, and its length is approximately 1.6 oral diam-
eters (0.6 total length). The diameter at the aboral end
where it joins with the bowl is 0.92 oral diameter. The sides
arc even, regular, not serrated or with anything more than
minor local bulges or contractions. Twelve turns of the spiral
lamina are included in this section of the collar; there are 1
or 2 small oval fenestrae in the seventh turn of the lamina.
The bowl is globular, and in general is evenly rounded, but
a little suggestion of flattening is obscurely apparent at the
aboral end. The bowl reaches its greatest diameter a little
below its middle, where it is 1.35 oral diameters (0.44 total
length). The bowl is 1.22 oral diameters or 0.33 total length
in length. The aboral end lacks any special differentiation
other than slight flattening.

The wall of the bowl is roughened by large, uneven-sized
blobs of tertiary alveoles, about 25 in number across one face.
Secondary prismatic structures occur between these blobs;
about 50 prisms, or more, are found across the middle of the
bowl. The wall of the collar is glassy-translucent, formed by
16 spiral turns, and under the highest powers of the micro-
scope faint primary prisms may be sometimes detected. The
small fenestrae that are included are about 0.17 oral diameter
in length, and the full width of the spiral turn across. The
edge of the spiral lamina is not thickened, nor does one turn
overlap the next; they simply join without special differentia-
tion. The wall of the bowl, in distinct contrast with that of
the collar, is dense and heavy. That of the collar is subuni-
formly about 0.0 1 oral diameter in thickness, whereas that ol
the bowl reaches nearly 0.1 at the equator. The junction of
collar and bowl is abrupt.

Length, total 88u, collar 53J.1; diameter, oral 33U, greatest
diameter of bowl 3811.

Aside from usual dimensional and proportional variations,
Codoncllopsis aleutiensis frequently has a less flaring suboral
region, a more or less cylindrical collar, a bowl proportion-
ately longer than wide, and the extraneous alveolar matter
replaced by a precise network of small secondary prisms.

Codoncllopsis aleutiensis is one of the medium-sized
species of the genus, and one of the few of northern distribu-
tion. It differs from the tropical C. ecaudata, and from close
relatives of that species, in the character of the bowl; in
ecaudata the bowl is smooth, trim, and lacking in blobs of
excess prismatic matter. The bowl of aleutiensis is more
rotund and obscurely flattened instead of having a tendency
toward pointing. The general facies of the whole lorica is
quite different from that of C. erythraensis, which species
also tends to have a flat-sided bowl. Codoncllopsis americana
has a proportionately longer bowl and the collar and bowl
are subequal, whereas in aleutiensis the bowl is much less

than half the length of the collar. Codoncllopsis aleutiensii
differs from C. oricntalis of the Japanese area in the rounded
instead of pointed bowl, distinctly longer collar, and more
slender build. Its coilar has a greater number of spiral turns
and the suboral funnel is better developed. It is not at all
like C. frigida, or any of the tropical species related to C.
orthoccras.

Recorded from ten stations in the Pacific, as follows: two
(117, 126) in the North Pacific middle latitudes, six (118,
in), 120, 121, 122, 123) in the East Asiatic marginal sea, and
two (124, 125) in the Alaskan secondary region. The distri-
bution of Codoncllopsis aleutiensis is exceptionally compact.

There are 17 pump and 12 net samples, of which 7 were
taken at the surface, 12 at 50 meters, anil 10 at 100 meters.
Maximum frequency, 125 loricae at station \zy, all other
records above minimum (2 to 100) except at station 126;
averages, 26.6 loricae and 22.5 per cent in pump and net
samples, respectively.

Temperature: pump samples 1 ?7i— 8?33 (4?qi), net
samples 2?oi-8?93 <4?95). Salinity: pump samples 32.73-
33.72 (33.02), net samples 32.76-34.06 (33.21). Density:
pump samples 25.47-26.52 (26.09), net samples 22.52-26.76
(26.22). pH: pump samples 7.84-8.04 (7.95), net samples
7.68-8.03 (7.82).

Type locality, station 120, at 100 meters; latitude 47 02'
north, longitude 166 20' east.

Codonellopsis americana Kofoid and Campbell

Codoncllopsis americana Kofoid and Campbell, 1929, p. 75, fig.
159.

The moderately tall lorica, with bowl and collar subequal,
slightly everted brim, cylindrical collar, rather elongated
rough bowl, and subrotund aboral end, has a length of 2.16
oral diameters. The oral margin is sharp-edged and has a
little trace of narrow brim. The cylindrical collar (0.41 total
length) has 8 spiral turns; of these the uppermost 3 are very
narrow, the middle 3 are twice as wide, and the last 2 are yet
wider. The fifth turn has a couple of faint oval fenestrae,
nearly the width of the turn. The diameter at the neck of
the collar is almost the same as the oral diameter; there is a
trace of lateral suboral contraction, and there is a slight
molding along the edges of each of the turns. The collar is
almost transparent. The dense, heavy bowl expands from
the neck to its greatest diameter of 1.54 oral diameters near
the equator. Below the equator it rapidly rounds off (82
to 120 ), and the aboral end is blunted.

The wall of the bowl reaches, in places, a thickness of 0.13
oral diameter. Its blobby, rough surface prevents accurate
determination. The exterior has large, irregular masses of
heavy alveolar tertiary structure.

Length, 76 to 112U.

The Carnegie loricae are shorter and stouter, and have
wider bowls and more pointed aboral ends than those from
the eastern tropical Pacific.

Codoncllopsis a man ana resembles C. indica (see Marshall,
1934, p. 639) closely, but the aboral end is less pointed, the
collar has traces of eversion, and it is without a wide group

24

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

of narrow suboral spiral turns; possibly these two species are
founded on extremes of a widely distributed species subject
to considerable physical variation.

Recorded from four stations, two each in the Atlantic and
the Pacific, as follows: one (2) in the Gulf Stream, one (33)
in the Caribbean Sea, one (40) in the Galapagos region, and
one (158) in the region of South Pacific island fields.

There are 2 pump samples (from the Caribbean Sea) and
} net samples, of which 1 was taken at the surface, 3 at 50
meters, and 1 at 100 meters. Frequency, minimum.

Temperature: Atlantic, pumpsamples28?25-28'; 49 (28^36),
net sample 20^35; Pacific, net samples i5?33-28?i4 (2i?73).
Salinity: Atlantic, pump samples 35.63-36.19 (35.90), net
sample 36.41; Pacific, net samples 34.89-35.58 (35.23). Den-
sity: Atlantic, pump samples 22.73-23.23 (22.98), net sample
25.76; Pacific, net samples 22.80-25.83 (24.31). pH: At-
lantic, pump samples 8.23-8.24 (8.235), net sample 8.21;
Pacific, net samples 7.87-8.39 (8.13).

Codonellopsis biedermanni (Brandt) Kofoid and Campbell
Codonellopsis biedermanni, Kofoid and Campbell, 1929, p. 75,

fig. 181.
The tall, rather slender lorica, with tightly wrapped spiral
collar, slim, elongated bowl, and pointed aboral horn, has a
length of 3.88 oral diameters. The thick oral margin is
entire. The collar flares (43°) within the upper one-sixth,
and then becomes virtually a cylinder with a diameter of
0.77 oral diameter for the remainder of its length (0.45 total
length). It is made up of 28 subequal spiral turns; the
uppermost 7 are, however, distinctly narrower than the
others. The heavy, narrow bowl forms a subcylindrical neck
(within 0.33 oral diameter), and then increases evenly to the
maximum diameter of 1.05 oral diameters at 0.7 total length
from the rim. Below this level it rapidly declines for about
0.74 oral diameter, reaching, at 0.12 total length above the
aboral end, only 0.36 oral diameter. The aboral horn con-
tracts below as a cone (33°). Its free end is slightly rounded,
its sides are full, and the very end is opened by a minute pore.

The wall of the collar is relatively thick (nearly 0.05 oral
diameter), and is paralleled by the wall of the bowl (0.065).
The horn is also thick (0.13). The wall of the collar has
thin, dark inner and outer laminae with three to four layers
of minute hexagons enclosed. The wall of the bowl has an
inner dark lamina, and there are two to four layers of
secondary hexagons or rectangles. The whole bowl is dense
brown, and the collar is a lighter shade. The outer surface
is neatly trim, and the bowl has small surface hexagons,
about 40 across its equator. The surface of the horn is retic-
ulated with larger areas. The horn has a narrow, sub-
cylindrical canal.

Length, 230 to 380^1.

Codonellopsis biedermanni is closest to C. meridionalis,
and differs from it in the more tapering, wider bowl as well
as in the less abrupt, less sharply distinct aboral horn: the
collar, also, is usually relatively longer. These are a unique
pair of species, hardly to be confused with others of the
group of longer species.

Recorded from two stations in the Pacific, as follows: one

(46) in the Galapagos region, and one (99) in the equa-
torial region.

There are 2 net samples, of which 1 was taken at 50 and

1 at 100 meters. Frequency, 3 per cent at station 99; average,

2 per cent.

Temperature, 23?2f5-27?84 (25^55); salinity, 35.04-35.33
(35.18); density, 22.50-24.13 (23.31); pH, 8.16-8.22 (8.19).

Codonellopsis brasiliensis (Brandt) Kofoid and Campbell
Codonellopsis brasiliensis, Kofoid and Campbell, 1929, p. 77,

fig- 153-
The short, trim lorica, with laterally concave cylindrical
collar, conical bowl, and pointed aboral end, has a length of
2.6 oral diameters. The oral margin is rounded over and
forms a minute brim. The generally cylindrical, hyaline
collar flares (20 ) in the upper 0.52 oral diameter and
reaches 0.59 oral diameter at the lower level. There are 6
spiral turns in this section. Below, it rapidly expands (14°)
to the neck, which has a diameter of 1.0 oral diameter and
includes 7 progressively wider spiral turns. Some local bulge
occurs near the upper end of the section, and one or two
round fenestrae are found in the tenth spiral turn. The
collar itself has a length of almost 0.47 total length. The
dense bowl is rotund; it expands from the neck to nearly 1.35
oral diameters at 0.65 total length, and rapidly (45 to 110°)
becomes convex conical below. The aboral end is sub-
hemispherically blunted.

The wall reaches 0.1 oral diameter in thickness across the
equator, and thins out above and below. The thickness of
the collar is about 0.02 oral diameter. There are large, sub-
rectangular tertiary prisms which enclose three to five layers
of smaller hexagons. The wall of the collar has thin laminae
which enclose a clear substance. The edges of the successive
spiral turns are not thickened. The surface of the bowl has
large, irregular areas with smaller hexagons between adjacent
patches.

Length, 76 to nip.

Codonellopsis brasiliensis bears close resemblance to C.
ccaudata, but has a distinct lateral concavity in the collar,
and large surface patches on the bowl; in this latter respect
it recalls C. americana and its relatives, but differs from
these in that the patches are never adherent blobs. Its collar
is clearly unlike that of C. turgescens, and its bowl is not
shaped like that of C. robusta.

Recorded from two stations (31, 33) in the Caribbean Sea,

in net samples taken at 100 meters. Frequency, 2 to 3 per cent.

Temperature, 22?56-23?i7 (22^86); salinity, 36.49-36.51

(36.50); density, 25.03-25.22 (25.12): pH, 8. 18-8. 19 (8.185).

Codonellopsis brevicaudata (Brandt) Kofoid and Campbell
(Figure 6)
Codonellopsis brevicaudata , Kofoid and Campbell, 1929, p. 77,

fig. 178; Marshall, 1934, p. 640.

The tall lorica, with greatly elongated collar with brim,

round, irregular bowl, and short aboral horn, has a length

of 5.1 oral diameters. The smooth oral rim forms a wide

(80°) brim. The distinctly elongated, hyaline, cylindrical

TAXONOMY AND DISTRIBUTION

25

collar continues with the same diameter (0.87 oral diameter)
to the howl, from which it is sharply set off, and is without
even traces of local modification. Its length is approximately
0.68 total length. The collar is made up of 26 spiral turns,
the uppermost 6 of which are progressively a little less wide
than the others: no fenestrae were found. The glohose.
dense howl, with a maximum diameter of 1.45 oral diam-
eters, arises from just below the collar, forming a round-
shouldered neck with a width of 0.14 oral diameter, ami
then rapidly curves to the equator. Below its equator the
howl rounds to the ahoral end. The aboral horn is conical
(25 ), and its free end is broken off (?) or else is normally
open.

The wall of the collar is rather thick (0.07 oral diameter)
and the spiral turns that compose it have optically dense
margins. The heavy bowl reaches a thickness little greater
than that of the collar (0.08). The collar has laminae which
enclose a hyaline substance, and the bowl has one to four
layers of small to large rectangular, hexagonal, or irregular
alveoles. The surface of the bowl is irregular, and some-
what hlobby near its equator, and has irregular secondary
structure. In the lorica under discussion there was a large
circular internal boss or button inside the mid-region of the
howl. The cavity of the horn is cut off from that of the
bowl by a thin diaphragm.

Length, 27511.

This lorica is more trim than those described elsewhere.
Marshall states that hers was 17811 in length.

Codonellopsis brevicaudata bears little resemblance to
others of the longer species of the genus. Its markedly irregu-
lar, swollen bowl, short ( ?) open horn, and very long, regular
collar are characters that distinguish it. The species of the
C. orthoceras group generally have neat bowls and long
aboral horns.

Recorded from one station (18) in the Sargasso Sea, in a
net sample taken at 100 meters. Frequency, minimum.

Temperature, 20?32; salinity, 36.81; density, 26.07; pH,
8.21.

Codonellopsis contracta Kofoid and Campbell
Codonellopsis contracta Kofoid and Campbell, 1929. p. 78, fig.

147-
The tiny, stout, ovate lorica, with collar shorter than bowl.
wide bowl, and broad, rounded aboral end, has a length of
2.5 oral diameters. The simple oral margin is thin-edged.
The collar increases in diameter from the margin to the
lower end, being a segment of a distinctly concave cone
(25 ) the length of which is less than 0.33 total length. It
has approximately 12 spiral turns, which progressively in-
crease in width as they descend. The lowermost of these
distinct, thin-edged turns has a single row of rounded
fenestrae (8 to 10) which are the full width of the turn.
The aboral flare of the collar merges into the bowl below.
The plump little bowl expands from the throat and reaches
2.0 oral diameters at its equator. Below the equator it
rapidly rounds off. The aboral end is hemispherical.

The wall of the bowl is thin (as nearly as could be deter-
mined, about 0.02 oral diameter). There are minule sec-

ondary (?) alveoles. The aboral half of the bowl is com-
monly plastered with lecal matter, ami small coccoliths are
frequent.

Length, 42 to 4941.

The tiny loricae of Codonellopsis contracta are difficult
to study, but the material collected appears to agree closely
with the typical forms. Length is probably related to tem-
perature.

Codonellopsis contracta is shorter (42 to 49(1) than C.
pusilla (56 to 59|.i)> and stouter (length 2.5 as against 3.08
oral diameters). Its bowl is wider and its aboral end broadly
rounded. The bowl lacks surface flecks, is never irregular,
and often has coccoliths. It is not closely related to C. ovata
or to C. inornata, both of which are also among the smaller
species.

Recorded from six stations, three each in the Atlantic and
the Pacific, as follows: two (3, 5-6) in the Atlantic drift,
one (13) in the Gulf Stream, one (123) in the East Asiatic
marginal sea, and one (130) in the California region.

There are 3 pump and 5 net samples, of which 3 were
taken at the surface, 2 at 50 meters, and 3 at 100 meters.
Maximum frequency, 16 per cent at station 130; other
records above minimum (4 to 8 per cent) from stations 5-6,
13; averages, 1.0 and 3.5 per cent in Atlantic net and Pacific
pump samples, respectively.

Temperature: Atlantic, net samples n?27-i3?79 (i2?58);
Pacific, pump samples 5?o-n?88 (8?6i), net sample 8?g6.
Salinity: Atlantic, net samples 32.68-36.06 (34.88); Pacific,
pump samples 32.87-33.72 (33.22), net sample 33.72. Den-
sity: Atlantic, net samples 24.94-26.95 (25.94); Pacific, pump
samples 25.13-26.14 (25.82), net sample 26.14. pH: At-
lantic, net samples 8. 10-8. 15 (8.12); Pacific, pump samples
7.96-8.1 1 (8.04), net sample 8.06.

Codonellopsis ecaudata (Brandt) Kofoid and Campbell
(Figure 7)

Codonellopsis ecaudata, Kofoid and Campbell, 1929, p. 79, fig.
154-

The short, neat lorica, with tall, cylindrical collar and
oval bowl with aborally rounded end, has a length of 2.88
oral diameters. The thin oral rim is sharp-edged. The
long, distinctly hyaline collar (0.47 total length) is a cylinder
with only the slightest traces of strictly local contractions and
bulges. It is made up of 13 spiral turns. Of these, the first
and third are very narrow, but the remainder are subuni-
form in width. There are 1 or 2 small, faint oval fenestrae
(the full width of a single turn) in the seventh and ninth
turns. The sides of the collar are not serrate, each turn
simply fitting on the top of the succeeding one with a fine,
dark external ridge along the edge as a sort of molding. The
oval, very dense bowl (contrasting sharply with the collar)
expands from the neck, which has the same diameter as the
oral opening, to a diameter of 1.47 oral diameters at 0.7 total
length below the oral rim. Below this widest level it con-
tracts rapidly, although with full convex sides of approxi-
mately 70 , to the aboral end. The aboral end is bluntly
but narrowly contracted or, less often, simply rounded off.

26

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

The wall of the bowl reaches almost 0.09 oral diameter in
thickness at the equator; it thins out above and below. That
of the collar is uniformly less than 0.03 oral diameter. There
are two or three (usually three) layers of thin-walled hex-
agonal prisms in the bowl between very thin inner and outer
hyaline laminae. In the collar the laminae enclose much
finer, primary alveoles. A faint tracery of tertiary structure
of coarser mesh encloses the surface mesh of secondary
hexagons.

Length, 98 to nou..

The lorica figured (fig. 7) has an uncommonly sharp
aboral end.

Codonellopsis ecaudata is close to C. robusta, C. brasiliensis,
and C. turgescens. From them it differs mainly in the shape
of the aboral end, which is flat in robusta, conical-pointed in
brasiliensis, and pointed in turgescens. The bowl is also flat-
sided in robusta, and wider in turgescens than in ecaudata.
The collar of turgescens is swollen in the middle, whereas
that of ecaudata is cylindrical. All these four species of the
equatorial sea are trim, and distinctly different from others.

Recorded from nine stations in the Pacific, as follows:
four (45, 46, 78, 80) in the Galapagos region, four (81, 82,
84, 85) in the region of South Pacific island fields, and one
(139) in the North Pacific trade region. Codonellopsis
ecaudata is, thus, fairly compactly distributed.

There are 7 pump and 8 net samples, of which 2 were
taken at the surface, 10 at 50 meters, and 3 at 100 meters.
Maximum frequency, 6 per cent at station 78; other records
above minimum (2 to 5 per cent) from stations 46, 80, 81;
averages, 3.1 loricae and 2.1 per cent in pump and net
samples, respectively.

Temperature: pump samples 22?78-26?42 (23:73), net
samples 22?37~27?89 (25?i5). Salinity: pump samples
35.18-36.17 (35.76), net samples 35.23-36.42 (35.79). Den-
sity: pump samples 23.56-25.11 (24.29), net samples 23.64-
24.33 ( 2 3-9 )- pH: pump samples 8.14-8.28 (8.18), net
samples 8.13-8.22 (8.17).

Codonellopsis inflata Kofoid and Campbell

Codonellopsis inflata Kofoid and Campbell, 1929, pp. 80-81,
fig. 176.

The moderately tall and stout lorica, with collar somewhat
longer than bowl, with everted rim, rotund bowl, and
slender, fairly long aboral horn, has a length of 3.74 oral
diameters. The oral rim is thin-edged and entire. The
collar (0.42 total length in length) has distinct suboral flare
(40 ) within the uppermost one-tenth. It is cylindrical,
with a diameter of 0.83 oral diameter at the aboral end, and
has a clean, unmodified lateral contour. There are about 38
decidedly wavy spiral turns, of which the anterior 18 are
very narrow (uppermost 0.37 of the collar), and the posterior
20 are subuniformly wider, yet as wavy, save for the last 2
or 3. The neck of the bowl has a width of approximately
0.29 oral diameter, and a diameter at its lower end of 1.0
oral diameter; its sides are rounded concave. The bowl ex-
pands from the lower end of the neck to 1.32 oral diameters
at 0.58 total length from the rim. Below this widest level it

rapidly contracts (90 ) to the aboral end, which has a diam-
eter of nearly 0.25 oral diameter. The aboral horn is almost
cylindrical, with a length of 1.0 oral diameter and with a
bluntly pointed free tip.

The wall of the collar hardly reaches 0.02 oral diameter in
thickness, but that of the more or less uniform bowl has a
thickness of 0.08. The collar is hyaline and the bowl is
dense. There are thin laminae in both collar and bowl; in
the layer thus enclosed are two or three rows of rounded
alveoles. The surface of the bowl has tiny, rounded sec-
ondary areas, as does also the horn. The horn has a wide
cavity, quite cut off from that of the bowl by a depressed
diaphragm.

Length, 130 to 22411..

The Carnegie loricae agree precisely with those of the
authors of the species.

Codonellopsis inflata has a wider bowl than any other of
the species related to C. orthoceras. Of these, it is closest to
C. pacifica, from which it may be distinguished by the
cylindrical aboral horn and the many wavy spiral turns in
the collar.

Recorded from one station (80) in the Galapagos region,
in a net sample taken at 50 meters. Frequency, 2 per cent.

Temperature, 26?o6; salinity, 35.95; density, 23.75; pH,
8.19.

Codonellopsis longa Kofoid and Campbell
(Figure 5)
Codonellopsis longa Kofoid and Campbell, 1929, p. 82, fig. 1S0.
Codonellopsis orthoceras. Hofker, 1931, pp. 367-369, figs. 47-
48A.

The elongated lorica, with long collar, egg-shaped bowl,
and long aboral horn, has a length of 4.35 oral diameters.
The oral margin is flat. The collar has little flare (53°),
well within the upper one-tenth. The long collar (0.45 total
length in length) is virtually a cylinder with a diameter at
the base of over 0.85 oral diameter. It is made up of 20
subequal spiral turns with molded edges. The bowl has a
length of 1.6 oral diameters and reaches its maximum
diameter of 1.21 oral diameters near 0.63 total length below
the rim. The neck is swollen to 1.0 oral diameter just below
its junction with the collar and contracted to 0.85 oral
diameter at 0.49 total length below the rim. The aboral
diameter of the bowl is only 0.28 oral diameter. The aboral
horn is conical (12 ), 0.82 oral diameter in length, and
sharply pointed.

The wall of the hyaline collar has a thickness of about
0.03 oral diameter, and that of the dense, brown bowl is
0.09 oral diameter. There are thin laminae which, in the
collar, enclose hyaline material; in the bowl there are two to
five layers of small hexagons, and larger tertiary structures
enclose the smaller hexagons in groups. The surface of the
bowl has small secondary hexagons enclosed by faint, larger
tertiary hexagonal meshwork. The cavity of the horn is
separated from that of the bowl by a thin diaphragm.

Length, 235 to 294^1.

Hofker's material may not all belong to Codonellopsis
longa; his figure 48A strongly suggests C. parva.

TAXONOMY AND DISTRIBUTION

Codonellopsis longa has a subcylindrical rather than wide
conical collar, the latter shape being distinctive of C. ortho-
ceras. It has a longer collar than C. pacifica, and, in addi-
tion, a longer bowl and horn than C. Speciosa. The collar
is concave in C. tessellata, and the bowl is egg-shaped rather
than potlike. It is not especially difficult to separate longa
from related species of the tropical plankton.

Recorded from thirty-six stations, thirteen in the Atlantic
and twenty-three in the Pacific, as follows: three (2, 15, id)
in the Gulf Stream, two (17, 18) in the Sargasso Sea, six
(22, 25, 27, 28, 29, 30) in the Atlantic equatorial region,
two (31, 32) in the Caribbean Sea, two (35-36, 99) in the
Pacific equatorial region, five (46, 47, 75, 77. 80) in the
Galapagos region, seven (48, 81, 82, 85, 95, 158, 159) in the
region of South Pacific island fields, three (100, 150, 151) in
the North Pacific trade region, one (113) in the North
Pacific middle latitudes, and five (133, 136, 137, 147, 149) in
the California region. Codonellopsis longa is a widely dis-
tributed species of the warmer regions of the ocean, and
avoids, as these and other records show, only the cooler
waters of the polar seas and waters contributed by them.

There are 8 pump and 42 net samples, of which 9 were
taken at the surface, 18 at 50 meters, and 23 at too meters.
Maximum frequency, 29 per cent at station 81; other records
above minimum (2 to 18 per cent) from stations 2, 16, 22,
32, 35-36, 46, 48, 75, 77, 80, 81, 85, 95, 99, too, 113, 136, 137,
147, 149, 151, 158, 159; averages in net samples, 2.9 and 6.8
per cent in the Atlantic and Pacific, respectively.

Temperature: Atlantic, pump samples 1 9 : 27-23? 64 ( 2 1 . 50 ) ,
net samples i 4 ?6o-2 7 ?88 (ai?57); Pacific, i8?5t-2 7 ?62
(24^35) and i8?4o-28?74 (24?32>, respectively. Salinity:
Atlantic, pump samples 36.41-36.47 (36.44), net samples
35.70-36.82 (36.37); Pacific, 34.70-36.46 (35.67) and 34.42-
36.44 (35.34), respectively. Density: Atlantic, pump samples
24.84-26.12 (25.48), net samples 23.26-26.62 (24.79); Pacific,
22.31-26.65 (24.35) and 22.31-25.55 (22.71), respectively.
pH: Atlantic, pump samples 8.21-8.23 (8.22), net samples
7.93-8.30 (8.19); Pacific, 8.18-8.31 (8.21) and 8.10-8.39
(8.25), respectively.

Codonellopsis meridionalis Kofoid and Campbell
Codonellopsis meridionalis Kofoid and Campbell, 1929, pp. 82-
83, fig. 177.

The stout lorica, with subequal collar and bowl and long
aboral horn, has a length of 3.82 oral diameters. The rather
thick oral margin is smooth and flat. The long collar (0.43
total length) is abruptly everted (50 ) well within the upper
one-tenth and forms an inverted truncate cone (5 ) in the
remainder. The diameter at its lower end is 0.83 oral diam-
eter. There are no local changes in lateral contour save as
the spiral turns form a serrate edge as they overlap one an-
other. There are 22 spiral turns, of which the anterior 10
are very narrow and the posterior 12 are at least twice as
wide. The wider turns are not strictly subequal in width.
The bowl forms a low neck, the width of which is approxi-
mately 0.33 oral diameter and the aboral diameter 1.0 oral
diameter. The rather stout bowl expands rapidly below the

truncate laterally concavc-subconical (22 ) neck, which is
rounded at its junction with the collar, and reaches a maxi-
mum diameter of 1.25 oral diameters at 0.59 total length
from the rim. The aboral part is rotund subangular (90°),
and reaches a diameter of 0.33 oral diameter near 0.8 total
length from the rim. The aboral horn is long (0.6 oral
diameter), conical (12 ), rough-walled, and blunt at the
free tip.

The wall of the collar reaches the unusual thickness of
nearly 0.05 oral diameter, and that of the bowl is nearly 0.16.
The wall of the collar has a dark inner lamina and there are
three to five layers of minute hexagons. The wall of the
bowl also has a thick inner lamina and two to four layers of
large secondary hexagons. The surface of the bowl is rather
trim, with small hexagons, but the horn is roughly canalicu-
late. The horn is penetrated by a wide canal and cut oil
from the cavity of the bowl by a thick diaphragm. Collar
and bowl are of a rich brown color, the latter more strongly

so.

Length, 197 to 245P.

Codonellopsis meridionalis is close to C. biedermanni,

from which species it differs in the abruptly set-off, narrow

aboral horn, tapering collar, and wider bowl. These two

species can scarcely be confused with others.

Recorded from three stations (35, 35-36, 155) in the

Pacific, all in the equatorial region.

There are 4 net samples, of which 1 was taken at the

surface, 2 at 50 meters, and 1 at 100 meters. Frequency, 50

per cent at station 35-36, 10 per cent at station 35; elsewhere,

minimum.
Temperature, 14:33-27:71 (i9°44)'> salinity, 34.88-34.93

(34.90); density, 22.46-26.06 (24.70); pH, 7.88-8.30 (8.03).

Codonellopsis minor (Brandt) Kofoid and Campbell
Codonellopsis minor, Kofoid and Campbell, 1929, p. 83, fig. 168.

The rather short lorica, with conical, everted collar, long
bowl, and short, conical aboral horn, has a length of 2.3 oral
diameters. The oral margin is sharp-edged. The collar
flares (32 ) within the upper one-tenth and is inverted sub-
conical (io°) for the remainder of its length. Its diameter
at the lower end is 0.92 oral diameter. In lateral contour it
is full but with local modifications. There are 1 1 spiral turns,
of which the anterior 2 are somewhat less wide than the
others. All the turns are outwardly concave and have
optically dark thickened margins. The neck of the bowl is
contracted from the throat, having a diameter at its lower
end of 0.85 oral diameter at 0.34 total length from the rim.
The line of junction with the collar and bowl is wavy. The
bowl rapidly expands below the neck to 1.25 oral diameters
at 0.57 total length below the rim. Below this equator it
contracts (85 ) as a decidedly convex hemisphere. The
aboral horn is short (0.33 oral diameter), conical (25 ), and
blunt; its anterior diameter is 0.26 oral diameter.

The wall of the collar is thick (over 0.08 oral diameter)
and that of the bowl is similar. There are thin laminae with
enclosed hyaline material in the collar, and two to three
layers of small hexagons in the bowl; in the bowl are also

28

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

larger, rectangular, thick-walled tertiary areas. The surface
of the bowl clearly shows the tertiary areas as hexagons,
about 20 in number, across its equator, and 10 to 14 second-
ary hexagons are enclosed by each of the thicker-walled,
larger areas. The horn is similar to the bowl. The cavity of
the horn is cut off from that of the bowl by a diaphragm
formed by the inner lamina. This cavity is wide and rather
closely follows the outer contour.

Length, 125 to 207(1.

Codonellopsis turgida and C. pura both have much shorter
and wider bowls than the present species. The collar of C.
minor is longer than in these two species. Codonellopsis
parva has a longer horn, wider bowl, and shorter, more
conical collar. Codonellopsis calijorniensis hardly has a horn
at all, and its bowl is wider than in minor. Codonellopsis
speciosa has a longer horn and different bowl.

Recorded from ten stations, five each in the Atlantic and
the Pacific, as follows: one (16) in the Gulf Stream, three
(24, 25, 27) in the Atlantic equatorial region, one (33) in
the Caribbean Sea, three (41, 46, 80) in the Galapagos
region, one (107) in the North Pacific trade region, and one
(153) in the Pacific equatorial region.

There are 9 pump and 3 net samples, of which 7 were
taken at 50 meters and 5 at 100 meters. Maximum fre-
quency, 3 per cent at station 46; frequency 2 per cent at
stations 16, 24; other records, minimum; average in Atlantic
net samples, 1.5 per cent.

Temperature: Atlantic, pump samples i4?6o-28?25(23?i3),
net samples 23?i2-23?64 (23?38); Pacific, 22?73-26?92
(24'; 12) and i4?55, respectively. Salinity: Atlantic, pump
samples 35.70-36.25 (36.11), net samples 36.00-36.41 (36.20);
Pacific, 34.73-36.21 (35.27) and 35.02, respectively. Density:
Atlantic, pump samples 23.23-26.02 (24.66), net samples
24.67-24.84 (24.75); Pacific, 22.63-24.70 (23.82) and 26.11,
respectively. pH: Atlantic, pump samples 7.93-8.30 (8.16),
net samples 8.14-8.23 (8.18); Pacific, 8.16-8.28 (8.21) and
7.92, respectively.

Codonellopsis orientalis Hada

Codonellopsis orientalis Hada, 19326, pp. 563-564, fig. 15.

The fairly tall, clean lorica, with subcylindrical collar,
ovate bowl, and broadly convex-conical aboral region, has a
length of 2.60 to 2.76 oral diameters. The thin oral margin
is rounded. The collar is generally cylindrical, although
there is slight suboral flare (45°) within the upper 0.21 of
the length of the collar in some loricae; in these the aboral
diameter of the collar is 0.91 oral diameter. The collar is
made up of 8 to 12 spiral turns, the uppermost 3 or 4 being
much narrower than the others, which are more or less uni-
form. Oval fenestrae (the full width of a turn) occur in the
sixth and also sometimes in the seventh turn. The collar has
a length of 0.3 to 0.4 total length, and commonly, but not
always, bulges in the middle (0.92 oral diameter). The
ovate bowl expands from the neck to its greatest diameter of
1.52 to 1.57 oral diameters near 0.55 total length from the
rim. Below this level the bowl contracts with full convex-
conical sides (85 to 105 ) to the blunt, rounded aboral end.

The wall of the collar is hyaline and uniformly thin (0.04

oral diameter), and has fine primary alveoles enclosed within
its laminae. The wall of the dense bowl is thicker (0.08
oral diameter) across the middle and thins above and below;
it is made up of large, blobby secondary prisms. The outer
surface is decidedly irregular, with coarse blocks of alveolar
material of diverse shapes.

Length, 84 to 931.1.

Codonellopsis orientalis bears some likeness to members
of the C. morchella group, but differs from them mainly in
the shape of the aboral end. This tropical assembly of
species has a distinctive fades rather difficult to describe in
words.

Recorded from one station (117) in the North Pacific
middle latitudes, in 2 net samples, 1 each taken at 50 and
100 meters. Frequency not over 2 per cent.

Temperature, 8?93-i2?56 (io?74); salinity, 34.06-34.22
(34.14); density, 25.89-26.41 (26.15); pH, 7.98-8.06 (8.02).

Codonellopsis pacifica ( Brandt) emended Kofoid
and Campbell

Codonellopsis pacifica, Kofoid and Campbell, 1929, p. 85, fig.
'75-

The moderately tall lorica, with collar longer than bowl,
wide, globose bowl, and long, narrow horn, has a length of
3.67 oral diameters. The thin oral margin is everted and
flaring (63 ) within the uppermost one-tenth. The collar is
0.41 total length, truncate, inverted conical (5 ), with a
diameter at the lowermost end of nearly 0.75 oral diameter,
and with bare lateral concavity. There are 24 spiral turns,
the upper 6 being narrow and the remainder wider. Of the
wide ones, some are narrower than others. The bowl has a
more or less cylindrical neck with a width of 0.3 oral diam-
eter, and a diameter at its lower end of 0.91 oral diameter.
The bowl rounds from this neck to 1.17 oral diameters at
0.57 total length below the rim, and then contracts convex-
conically (90 ) to the aboral end, the diameter of which is
less than 0.33 oral diameter. The conical (6°) aboral horn
has a length of 1.0 oral diameter, and its free tip is blunt.

The wall of the collar is thin (0.02 oral diameter), and that
of the bowl is thicker (0.08). Thin, hyaline laminae enclose
colorless material, and the bowl has single or double layers
of large, often rectangular secondary prisms. The surface
of the bowl has rather small rounded areas, and scattered yet
larger round areas of several sizes. Coccoliths are common
on the surface. The horn has a similar surface and is hol-
low and irregularly but widely canaliculate, and has a thin,
depressed diaphragm which cuts its cavity from that of the
bowl. The collar is glass-clear and the bowl dense.

Length, 195 to 270^1,

Codonellopsis pacifica differs from C. longa in being
shorter and rather more plump, and having a shorter horn
and a collar with fewer turns. The bowl is not so expanded
as that of C. inflata; the collar is longer than in C. speciosa,
and less conical than in C. orthoceras.

Recorded from four stations in the Pacific, as follows:
three (45, 46, 78) in the Galapagos region, and one (81) in
the region of South Pacific island fields.

TAXONOMY AND DISTRIBUTION

29

There are 5 net samples, all taken at 50 meters. Fre-
quency, 2 per cent at stations 78, 81.

Temperature, 22:37-26=42 (23^94); salinity, 35.23-36.03
(35.55); density, 23.56-24.33 (24.08); pH, 8.13-8. 17 (8.15).

Codonellopsis parva Kofoid and Campbell

(Figures 2, 3)
Codonellopsis parva Kofoid and Campbell, 1929, p. 86, fig. 170.
The rather short, stout lorica, with collar and bowl more
or less subequal, and with short neck, rounded bowl, and
conical aboral horn, has a length of 2.16 to 3.34 oral diam-
eters. The oral margin is thin-edged, and there is some
suboral eversion with flare (58°), or even a flat brim. The
collar (0.32 to 0.38 total length) is a segment of an inverted
truncated cone (3 to 4 ) with a diameter at the aboral end
of 0.79 oral diameter. The sides show almost no lateral
modification in contour. The collar is made up of 15 to 17
spiral turns, of which the anterior 3 or 4 are a little narrower
than the others; the remainder are subequal, and all have
thin, molded edges. One or two of the turns overlap. The
neck is rounded, with a maximum diameter of 0.93 oral
diameter and contraction below (0.83 oral diameter); this
contraction is at 0.37 to 0.46 total length below the rim and
marks the lower limit of the neck. The bowl swells rapidly
from the neck and reaches as much as 1.32 oral diameters
near 0.56 total length. The full, convex lower bowl con-
tracts evenly and reaches 0.28 oral diameter at 0.83 total
length. The aboral horn is conical (n° to 15°), with a
length of 0.52 to 0.62 oral diameter, and has a blunted to
sharply pointed free tip.

The wall of the collar is only 0.02 oral diameter in thick-
ness, or less, but that of the bowl is nearly 0.07. The bowl
has large, mostly rectangular tertiary areas in one or two
layers, with pallid contents. The surface of the bowl has a
subequal mesh of tertiary areas; rarely are there (4) hemi-
spherical bosses of small secondary hexagons built into the
equatorial region. The wall of the horn is reticulate. The
cavity of the bowl is separated from that of the horn by a
thin diaphragm.

Length, 145 to 1871.1.

The Carnegie loricae are longer and commonly more
slender than those described elsewhere.

Codonellopsis parva has a shorter, less stout, narrower
bowl and longer horn than C. pura. Its collar is not so long
as that of C. speciosa, and its horn is narrower and more
abruptly limited. The horn is not so long as in C. minor,
and the bowl not so swollen as that of C. turgida. Codo-
nellopsis pacifica has a taller collar and much longer horn.
Recorded from thirteen stations, seven in the Atlantic and
six in the Pacific, as follows: one (2) in the Gulf Stream,
one (20) in the Sargasso Sea, three (23, 25, 28) in the
Atlantic equatorial region, two (33, 34) in the Caribbean
Sea, one (77) in the Galapagos region, one (100) in the
North Pacific trade region, three (153, 154, 155) in the
Pacific equatorial region, and one (157) in the region of
South Pacific island fields.

There are 15 net samples, of which 2 were taken at the

surface, 6 at 50 meters, and 7 at 100 meters. Maximum
frequency, 26 per cent at station 154; other records above
minimum ( 2 to 8 per cent) from stations 2, 20, 23, 25, 77,
100. 153, 157; average in the Pacific, 8.4 per cent.

Temperature: Atlantic, l4?6o-26?79 (22?i2); Pacific,
22?73-27?76 (25?96). Salinity: Atlantic, 35.70-36.73 (3 6 -3 2 >:
Pacific, 34.71-36.04 (35.08). Density: Atlantic, 23.79-26.02
(25.18); Pacific, 22.31-24.53 (23.16). pH: Atlantic, 7.93-
S.26 (8.16); Pacific, 7.93-8.32 (8.20).

Codonellopsis pura (Brandt) emended Kofoid and Campbell
(Figure 1)

Codonellopsis pura, Kofoid and Campbell, 1929, pp. 86-87, fig.

169.
The short, plump lorica, with collar shorter than bowl,
wide bowl with stout neck, and tiny knoblike aboral horn,
has a length of 2.22 oral diameters. The thin oral margin
has a recurved brim (not clearly shown in the figure), and a
subcylindrical collar the length of which is only 0.29 total
length; its aboral diameter is 0.83 oral diameter. Local
modifications are absent. There are 9 subequal spiral turns,
each with thickened edges. The neck is a convex ring. Its
greatest diameter is 0.89 oral diameter, its least, at the lower
end, is 0.83, and its width is nearly 0.29 oral diameter. The
rotund bowl swells out to a maximum, at its equator, of 1.29
oral diameters; this is at 0.63 total length from the rim.
Below its equator the bowl is hemispherical. The short
aboral horn is conical (25°), and only 0.2 oral diameter in
length; its free lip is squared off.

The wall of the collar is only 0.02 oral diameter in thick-
ness, and that of the bowl is subuniformly nearly 0.07. There
is a dark inner lamina. The bowl is rather pallid, with only
large, narrow, rectangular tertiary areas clearly marked. The
surface shows large, thin-walled, overlapping circles. The
collar is nearly transparent and the bowl is somewhat denser.
A diaphragm separates the cavity of bowl and horn.
Length, 125 to 195(1.

The Carnegie loricae are shorter, with a convex neck, and
have shorter horns than the originals.

Codonellopsis pura has a shorter horn, a wider bowl, and
a shorter collar than either C. parva or C. minor. Its collar
is shorter and its horn better developed than in C. ealijorm-
ensis. The bowl is not so wide as that of C. turgida.

Recorded from eleven stations, six in the Atlantic and five
in the Pacific, as follows: one (18) in the Sargasso Sea, five
(22, 23, 24, 25, 27) in the Atlantic equatorial region, two
(77, 80) in the Galapagos region, two (81, 157) in the region
of South Pacific island fields, and one (100) in the North
Pacific trade region.

There are 2 pump samples from the Pacific, and 10 net
samples, of which 6 were taken at 50 meters and 6 at 100
meters. Maximum frequency, 6 per cent at station 81; fre-
quency 4 per cent at station 157; all the remainder mini-
mum; the average frequency in Pacific net samples is 3.6
per cent.

Temperature: Atlantic, net samples i4?6o-26?04 (i9?i6);
Pacific, pump samples 23?69~27?62 (25?65), net samples

3°

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

26?o6-27?03 (26?47). Salinity. Atlantic, net samples 35.61—
36.81 (36.07); Pacific, pump samples 34.70-36.04 (35.37),
net samples 35.24-35.95 (35.68). Density: Atlantic, net
samples 23.98-26.62 (25.66); Pacific, pump samples 22.31-
24. 54 (23.42), net samples 22.91-23.75 (23.40). pH: At-
lantic, net samples 7.93-8.30 (8.13); Pacific, pump samples
8.19-8.22 (8.20), net samples 8.19-8.32 (8.23).

Codonellopsis pusilla (Cleve) Kofoid and Campbell

(Figure 8)

Codonellopsis pusilla, Kofoid and Campbell, 1929, p. 87, fig.
146; Hada, 19326, pp. 561-562, fig. 12.

The short, plump lorica, with short collar of few turns and
longer swollen bowl with blunted aboral end, has a length of
3.08 oral diameters. The oral margin is thin and rounded.
The hyaline collar is a concave segment of a cone (14 ) the
basal diameter of which is 1.28 oral diameters. It is com-
posed of 5 turns, the lower 3 of which are wider than the
upper 2. There is a single ovate fenestra between the last 2
turns. The swollen, dense bowl expands from the throat,
reaches nearly 2.0 oral diameters at its equator, and then
contracts (within 125°) to the broadly oval aboral end.

The wall reaches 0.07 oral diameter in thickness in the
bowl, where it is practically uniform; in the collar it is less
than half as thick. There is only a single layer of rectangular
secondary alveoles in the bowl; in both collar and bowl very
fine primary alveoles are visible under the best magnifica-
tion. The outer surface is made up of distinct but small
hexagons. Rarely are there a few agglomerated alveolar
flecks on the surface.

Length, 56 to 59^1.

Some loricae have pointed instead of ovate aboral ends.
The collar may have as many as 13 narrow spiral turns,
fenestrae may be lacking, and the length may reach 2.6 oral
diameters.

Codonellopsis pusilla differs from C. contractu in size and
proportions, in having fewer fenestrae if any, and in the
shape of the aboral end, which is decidedly broad in con-
tracta. These two are the smaller species of the cooler,
northern ocean, and are not readily comparable with those
of the tropics or Antarctic; among the latter are two (C.
glacilis and C. gaussi) peculiar species, the origin of which
must have been in tropical species. So also must have been
the origin of the northern neritic C. jrigida and C. borealis.
The former of these last two species is much like pusilla.

Recorded from five stations, two in the Atlantic and three
in the Pacific, as follows: two (5-6, 6) in the Atlantic drift,
two (116, 117) in the North Pacific middle latitudes, and
one (118) in the East Asiatic marginal sea.

There are 1 pump sample and 6 net samples, of which 3
were taken at the surface, 3 at 50 meters, and 1 at 100
meters. Maximum frequency, 50 per cent at station 117;
other records above minimum (2 to 23 per cent) from
stations 6, 116, 118; averages, 11.2 and 33 per cent in Atlantic
and Pacific net samples, respectively.

Temperature: Atlantic, net samples n?62-i2?44 (n?97);
Pacific, pump sample io?i8, net samples n?i8-i2?56

(n?87). Salinity: Atlantic, net samples 35.51-35.55 (35-53);
Pacific, pump sample 33.61, net samples 33.79-34.22 (34.00).
Density: Atlantic, net samples 26.95-27.21 (27.03); Pacific,
pump sample 25.85, net samples 25.83-25.89 (25.86). pH:
Atlantic, net samples 8.08-8.15 (8.12); Pacific, pump sample
8.21, net samples 8.06-8.1 1 (8.08).

Codonellopsis speciosa Kofoid and Campbell
(Figure 4)
Codonellopsis speciosa Kofoid and Campbell, 1929, p. 88, fig. 174.

The elongated lorica, with long, little-everted collar, stout
bowl, and stout, long aboral horn, has a length of 4.2 oral
diameters. The oral margin is thin-edged and sharp. The
long collar (0.42 total length) has some suboral eversion and
a low, rounded brim. The collar is a truncated segment of
an inverted cone (8°) and has a basal diameter of 0.81 oral
diameter. It is made up of 18 subequal spiral turns. The
neck is swollen (0.85 oral diameter) and then constricted
(0.81 oral diameter), all within 0.22 oral diameter. The
wide, elongated bowl rapidly swells to a maximum of 1.26
oral diameters at 0.59 total length below the rim. Below
this level it gradually decreases (75 to 80°), reaching at
0.75 total length a diameter of nearly 0.56 oral diameter.
Below this last level it becomes a short segment of an in-
verted truncated cone (35°) with a length of 0.37 oral diam-
eter, and has a diameter at its lower end of almost 0.3 oral
diameter. The aboral horn is conical (15°), with a length
of 0.7 oral diameter, and is blunted at its free tip.

The thin, hyaline collar (0.02 oral diameter in thickness)
is in contrast with the thicker (0.075), dense bowl. There
are thin laminae which in the collar develop hyaline enclosed
material, and in the bowl two or three layers of small prisms;
large tertiary structures in one or two layers enclose the
secondary prisms. The surface of the bowl shows a faint
tracery of large, circular tertiary areas. The cavity of the
bowl is cut off from that of the horn by a horizontal dia-
phragm.

Length, 181 to 225^1.

In the aboral projection of the bowl as well as in the
greater elongation, the Carnegie loricae differ from those
originally described. The collars are longer, and the horns
more emergent.

Codonellopsis speciosa has a longer, stouter bowl and
shorter, stouter horn than C. pacified. Its collar is longer, its
bowl different, and its horn longer than in C. part/a. Codo-
nellopsis inflata has an even wider, shorter bowl, a longer,
more slender aboral horn, and a collar with wavy margins
on the spiral turns.

Recorded from three stations, one in the Atlantic anil two
in the Pacific, as follows: one (23) in the Atlantic equatorial
region, one (45) in the Galapagos region, and one (152) in
the Pacific equatorial region.

There are 2 pump samples from the Pacific, ami 3 net
samples, of which 2 were taken at 50 meters and 3 at 100
meters. Maximum frequency, 2 per cent at station 152;
other records all minimum; average in Pacific pump samples,
1.5 loricae.

TAXONOMY AND DISTRIBUTION

3 1

Temperature: Atlantic, net sample 3&?02; Pacific, pump
samples n?4&-i4?32 (i2?9o), net samples 2i?69~22?37
I 22. 115). Salinity: Pacific, pump samples 34.50-34.73 (34.61),
net samples 35.21-35.23 (35.22). Density: Pacific, pump
samples 25.75-26.50 (26.12), net samples 24.30-24.48 (24.39).
pH: Atlantic, net sample 8.18; Pacific, pump samples 7.76-
7.87 (7.81), net samples 8. 12-8. 13 (8.125).

Codonellopsis turgescens Kofoid and Campbell
mettopsis turgescens Kofoidand Campbell, 1929, p. 90, fig.

'55-

The short, neat lorica, with swollen cylindrical collar,
rather wide, distinctly oval bowl, and pointed aboral end,
has a length of 3.5 oral diameters. The thin oral rim is
sharp-edged. The collar flares (37°) within its upper one-
tenth, expands slowly and evenly, and reaches 1.17 oral
diameters near 1.0 oral diameter below the rim. This swollen
region occupies a band with a width ot approximately 0.8
oral diameter. The diameter of the collar at the neck is
about 0.91 oral diameter. The collar itself has a length of
0.43 total length and there are 14 subuniform spiral turns.
The clean-cut bowl expands from the neck rapidly, reaching
1.43 oral diameters at 0.62 total length below the rim, and
rapidly contracts as an inverted full convex-conical (47 to
no°) section. The aboral end is barely pointed.

The wall reaches nearly 0.09 oral diameter in thickness at
the equator of the dense, brown bowl, and rapidly thins
above and below. That of the collar is uniformly about
0.03 oral diameter. There are two or three layers of thin-
walled, hexagonal prisms in the bowl, enclosed by laminae.
In the hyaline collar there are very fine alveoles. A faint
tertiary mesh encloses irregular groups of secondary meshes
in the bowl.

Length, 88 to ioou.

Codonellopsis turgescens resembles C. ecaudata, but differs
in having a swollen collar instead of a cylindrical one. From
C. robusta it differs in that the latter has a short, potlike
bowl as well as a different collar. Codonellopsis turgescens
may be distinguished from C. brasiliensis mainly by the
collar, which in brasiliensis is concave-cylindrical. The shape
of the aboral end is less distinctive in turgescens than in
robusta.

Recorded from one station (46) in the Galapagos region,
in a net sample taken at the surface. Frequency, minimum.

Temperature, 23?3o; salinity, 35.32; density, 24.11; pH,8.i6.

Codonellopsis turgida Kofoid and Campbell
(Figure 9)

Codonellopsis turgida Kofoid and Campbell, 1929, p. 90, fig. 167.

The short, stout lorica, with short, wide collar, very plump,
rotund bowl, and strictly conical aboral horn, has a length of
1.75 oral diameters. The thin oral margin is sharp-edged.
The collar (0.63 total length) is an inverted truncated cone
(18 ) with a lower diameter of 0.84 oral diameter; the full
sides lack even local modifications in contour. There are 12
subuniform, narrow spiral turns with dark edges and plane
sides. The bowl has a very low neck, the lower diameter of
which is only a little more than at the throat, and which is at

0.66 total length from the rim. The bowl rapidly swells to
1.24 oral diameters at 1.0 oral diameter from the rim. Below
its equator the bowl decreases as a hemisphere in diameter.
The aboral horn (0.18 oral di. muter in length) is conical
(45 ), with a basal diameter oi [.33 its own length, and is
blunt at the free tip.

The wall of the collar is only 0.02 oral diameter in thick-
ness, and that of the bowl is thicker (0.04). There are thin
laminae which in the collar enclose hyaline material, but
which in the bowl enclose two or three layers of hexagons.
The surface has uniform small circles of secondary prisms.
The contour of the dense bowl is irregular. The cavity of
the horn is cut off from that of the bowl by a continuation of
the inner lamina of the bowl.

Length, 1 12U.

The Carnegie loricae are shorter (112U. as against 143 to
170U.) and wider and have more and narrower spiral turns
in the collar than do those figured elsewhere, and have no
suboral flare.

The wide, swollen bowl and short horn at once set Codo-
nellopsis turgida apart from its near relatives, C. pura and
C. parva. Its bowl is not so long as in C. minor, nor is the
horn so short as in C. calif orniensis.

Recorded from five stations, two in the Atlantic and three
in the Pacific, as follows: two (23, 25) in the Atlantic equa-
torial region, one (77) in the Galapagos region, one (81) in
the region of South Pacific island fields, and one (99) in the
Pacific equatorial region.

There are 5 net samples, of which 1 was taken at the
surface, 1 at 50 meters, and 3 at 100 meters. Maximum fre-
quency, 9 per cent at station 99; frequency, 4 per cent at
station 81; other records minimum; average in the Pacific,
4.6 per cent.

Temperature: Atlantic, 14^60; Pacific, 22?72-27?84
(24?29). Salinity: Atlantic, 35.70-36.02 (35.86); Pacific,
35.04-36.04 (35.64). Density: Atlantic, 26.62; Pacific, 22.50-
24.53 (23.53). pH: Atlantic, 7.93-8.18 (8.05); Pacific, 8.19-
8.22 (8.20).

DICTYOCYSTIDAE Haeckel emended
Dictyocystidae, Kofoid and Campbell, 1929, p. 285.

The family includes four genera: Luminella, Wangiella,
Dictyocysta, and Wailesia. Of these only Dictyocysta was
found in the Carnegie material. The other genera arc
coastal, and Luminella and Wailesia are being newly estab-
lished by Kofoid and Campbell (1939, p. 284). Wangulla is
also rather recently established (D. Nie, 1934). The family
is allied to the Codonellidae but not closely, and to the
Codonellopsidae but perhaps more distantly. Dictyocysta is
eupelagic.

DICTYOCYSTA Ehrenberg emended

Dictyocysta, Kofoid and Campbell, 1929, p. 285.

The extraordinary genus Dictyocysta is allied to Luminella.
a genus being established by Kofoid and Campbell (1939,
p. 284). Luminella is like Stenosemella save that the collar
has demilunes. Closer, perhaps, is Wangiella, with W. dicol-

¥

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

laria, from the coast of China. Kofoid and Campbell (1929)
placed Dictyocysta among the more complex Tintinnoina,
between the Undellidae and Tintinnidae, only to insure its
isolation from Codonella, to which it had previously been
allied.

Dictyocysta is found abundantly in the tropics, although
some species are northern in distribution in the Atlantic.
None is antarctic.

Seventeen species are described here.

Dictyocysta dilatata Brandt

Dictyocysta dilatata, Kofoid and Campbell, 1929, p. 288, fig. 549.

The regular, fairly large lorica, with mitra-Yikt fenestra-
tion, distinct set-off collar, and pointed aboral end, has a
length of 1.4 oral diameters. The oral rim has 8 flattened
arches and is decidedly undulating. The collar is inverted
subconical, flaring (22 ); its length is 0.23 total length; its
diameter at the lower end is 0.89 oral diameter; and it has 8
rounded, subequal, rectangular windows with heavy beams
and mullions. The bowl is acorn-like, without shoulder, and
reaches a maximum diameter of 1.0 oral diameter at 0.58
total length from the oral rim. The aboral region is convex
conical (62 ), and the aboral end is pointed.

The wall of the bowl has a maximum of 7 vertical rows
of unequal, squarish to rounded, rarely subpentagonal
fenestrae. These fenestrae number 12, 12, 16, 8, 6, and 5
respectively in horizontal rows. The smaller ones are mostly
in the aboral 3 rows. The framework which encloses the
windows is thick and heavy. All windows are closed by
hyaline panes.

Length, 58 to 70^1.

Dictyocysta dilatata is of the general D. mitra type. It
differs from mitra in the shape of the collar, pointed aboral
end, and trimmer form. Dictyocysta obtusa lacks clear dis-
tinction between collar and bowl, as does also D. jenestrata;
and D. spinosa, though of similar type, has oral spinules. On
the whole, dilatata is a clear-cut, easy species to recognize.

Recorded from ten stations in the Pacific, as follows: four
(54, 62-63, 64, 65) in the South Pacific middle latitudes, one
(68) in the Galapagos region, one (no) in the North Pacific
trade region, two (131, 148) in the California region, and
two (142, 145) in the North Pacific middle latitudes.

There are 4 pump and 8 net samples, of which 3 were
taken at the surface and 9 at 100 meters. Maximum fre-
quency, 12 per cent at station 131; other records above mini-
mum from stations 54, 65, 68, 145; average in net samples,
4.3 per cent.

Temperature: pump samples i6?58-20?io (i7?88), net
samples I2?i2-20?6r (i-]°.6i). Salinity: pump samples
34.47-34.95 (347 1 )' net samples 34.30-35.35 (34-5 6 )- Den-
sity: pump samples 24.72-25.15 (24.96), net samples 24.33-
25.52 (25.00). pH: pump samples 8.14-8.31 (8.24), net
samples 8.10-8.32 (8.15).

Dictyocysta duplex Brandt emended Kofoid and Campbell
Dictyocysta duplex, Kofoid and Campbell, 1929, p. 289, fig. 564.
The large lorica, with collar resembling a colonnade of a
Greek temple, globose bowl, and "duplex" fenestration, has

a length of 1.56 oral diameters. The oral rim is smooth, and
lacks the arches characteristic of many other species. The
collar is 0.75 oral diameter in length, cylindrical, and formed
by 7 tall, rectangular, subequal windows with rounded edges
(height of windows is 1.2 their width) and with hyaline,
rodlike beams and mullions. All the windows are open. The
bowl is globular, with its maximum diameter of 1.22 oral
diameters near its middle. There is no distinct shoulder, the
bowl simply rounding from the neck to its widest level. The
aboral end is hemispherical.

The wall of the bowl is made up of large doubled or
"duplex" overlapping circles with hyaline areas enclosed.
Overlapping areas commonly include smaller circles. The
circles are the sites of coccoliths.

Length, 60 to 75U..

Dictyocysta duplex bears some similarity to D. polygonata,
but the duplex wall and collar are different. None of the
other species has similar walls, and, indeed, this character
is shared only by isolated examples in other genera, espe-
cially Codonella and Codonaria.

Recorded from four stations, one in the Atlantic and three
in the Pacific, as follows: one (23) in the Atlantic equa-
torial region, one (48) in the region of South Pacific island
fields, one (100) in the North Pacific trade region, and one
( 148) in the California region.

There are 2 pump and 2 net samples, of which 2 were
taken at the surface and 1 each at 50 and 100 meters. Maxi-
mum frequency, 3 per cent at station 148; other records 2
per cent.

Temperature: Atlantic, net sample 20?90; Pacific, pump
samples 20?io-27?73 (23?9i), net sample 23763. Salinity:
Atlantic, net sample 36.04; Pacific, pump samples 34.72-34.95
(34.83), net sample 36.44. Density: Atlantic, net sample
25.30; Pacific, pump samples 22.29-24.72 (23.50), net sample
24.86. pH: Atlantic, net sample 8.14; Pacific, pump sample
8.21, net sample 8.23.

Dictyocysta elegans Ehrenberg emended Kofoid
and Campbell

Dictyocysta elegans, Kofoid and Campbell, 1929, pp. 289-290,
fig. 570.

The moderately large lorica, with double row of windows,
acorn-like bowl with 3 rows of fenestrae, and rounded aboral
end, has a length of 1.46 oral diameters. The undulating
oral margin has a thin vertical list and 8 flattened arches.
The collar is cylindrical, with a length of 0.5 total length,
and with the diameter at the lower end equal to that of the
oral margin. The upper half of the collar has 8 inverted
pentagonal windows, and the lower part has 10 smaller,
squarish to pentagonal ones. The beams and mullions are
heavy and rounded, and the windows are all open. The
bowl is shaped like an acorn cup with traces of shoulder
(1.1 oral diameters in diameter) a little below the lower
limit of the collar, and thence it rapidly contracts (53 in
the flat-sided upper two-thirds, and 8g° lower down). The
aboral end is rounded.

The bowl has 3 rows of major, closed fenestrae; one of
these rows, of 10 subequal, rounded fenestrae, is near the

TAXONOMY AND DISTRIBUTION

33

shoulder region; the second row, of 12 larger, more or less
circular ones, is equatorial; and the third row, of 6 large
circular ones, is postequatorial. In addition, there are scat-
tered smaller ones on the howl. The interfenestral area is
reticulated. Primary alveoles occur in the upper framework
and between the reticulations on the bowl.

Length, 70 to 77U.

Dictyocysla elegans differs from D. magna in shortness
and in less conical bowl, from D. inaequalis in less ovoid
howl, from D. ampla in less shoulder and in bowl shape, and
from D. speciosa in size and bowl shape. The bowl is not
so pointed as in D. miilleri and it has different proportions
from that of D. fundlandica. The remaining species differ
from elegans in having only a single row of windows.

Recorded from seven stations, two in the Atlantic and five
in the Pacific, as follows: one ( 14) in the Gulf Stream, one
(23) in the Atlantic equatorial region, three (116, 117, 128)
in the North Pacific middle latitudes, one (118) in the
East Asiatic marginal sea, and one ( 130) in the California
region.

There are 7 pump and 7 net samples, of which 1 was
taken at the surface, 9 at 50 meters, and 4 at 100 meters.
Maximum frequency, 5 per cent at station 130; other records
above minimum (2 to 4 per cent) from stations 116, 117;
average in net samples from the Pacific, 3 per cent.

Temperature: Atlantic, pump sample 14^95, net sample
20?99; Pacific, io?i8-i2?9i (n°44) and 8? 9 3-i2? 9 i
(io?79), respectively. Salinity: Atlantic, pump sample 35.10,
net sample 36.04; Pacific, 33.07-34.22 (33.53) and 33.14-34.22
(33.72), respectively. Density: Atlantic, pump sample 26.08,
net sample 25.30; Pacific, 25.13-25.89 (25.56) and 25.19-
26.41 (25.82), respectively. pH: Atlantic, pump sample 8.18,
net sample 8.14; Pacific, 8.00-8.26 (8.13) and 7.98-8.26
(8.08), respectively.

Dictyocysta inaequalis Kofoid and Campbell
Dictyocysta inaequalis Kofoid and Campbell, 1929, p. 293, fig.
57i-

The rather tall lorica, with elegans-\ike fenestration, full
ovoid bowl, and rounded aboral end, has a length of 1.6
oral diameters. The oral rim is undulating, with 7 flat
arches and a thin list. The collar is cylindrical, with lateral
concavity, with a length of 0.54 total length, with the least
diameter near its middle (0.89 oral diameter), and with 7
subpentagonal, subequal windows in the upper row and 9
quite irregular ones in the lower row. The bowl swells
regularly from the throat, reaches 1.43 oral diameters a little
below the neck, and then contracts convexly (93° in the
upper three-fifths and 105 in the aboral section). The
aboral end is rounded off evenly.

The wall of the bowl has 2 postequatorial rows of fenestrae,
of which the upper row of 12 are large circles, subequal, and
the lower row of 6 are unequal, less regular subcircles. A
few odd, scattered ones occur. The interfenestral surface is
reticulated with many small polygons. The windows of the
collar are apparently open, and the rather thick beams and
mullions are hyaline.

Length, 64 to 66u.

Dictyocysta inaequalis resembles D. ampla closely, but has
a less full bowl, less regular lower row of fenestrae, and
fainter polygons. Its bowl is more convex than that of D.
elegans, and its collar is laterally concave.

Recorded from one station (23) in the Atlantic equatorial
region, in a net sample taken at 50 meters. Frequency, mini-
mum.

Temperature, 20^99; salinity, 36.04; density, 25.30; pH,
S.14.

Dictyocysta lata Kofoid and Campbell

Dictyocysta lata Kofoid and Campbell, 1029. y. 293, litf- 562.

The rather short, squat lorica has tall, rectangular win-
dows in the collar, a wide, dishlike bowl, and /cpida-\ikc
fenestration, and its length is 1.18 oral diameters. The oral
margin undulates only slightly. The collar is cylindrical,
with a length of 0.44 total length, and has 8 tall, rectangular
windows. The beams and mullions are relatively thin. The
dishlike bowl is shaped much like a deep mush bowl. It
lacks shoulders, and contracts, or rather rounds, from the
throat to the aboral end, forming virtually a hemisphere.
The aboral end is unmodified.

The wall of the bowl has a single irregular equatorial row
of 8 very large, roughly ovoid fenestrae. The remaining
surface is faintly reticulated, and the framework of the
collar glistens. The collar windows are open.

Length, 57 to 62|i.

Dictyocysta lata differs from D. reticulata not only in the
shape of the bowl, but in the size and irregularity of the
fenestrae. It has a shorter, fuller bowl than does D. grandis,
and is much shorter. It has only a single row of fenestrae,
and a bowl differently shaped from that of D. lepida. The
bowl lacks the shoulder and distal contraction of D. mexicana
and is less dense than in most species.

Recorded from one station (27) in the Atlantic equatorial
region, in a net sample taken at 100 meters. Frequency,
minimum.

Temperature, i8?o8; salinity, 36.03; density, 26.06; pH,
8.09.

Dictyocysta lepida Ehrenberg emended Kofoid and Campbell

Dictyocysta lepida, Kofoid and Campbell, 1929, p. 294, fig. 558;
Alzamora, 1929, p. 11, fig. 25.

The moderately small lorica, with single row of collar win-
dows and generally acorn-shaped bowl with 2 minor and 1
major rows of fenestrae, has a length of 1.28 oral diameters.
The oral margin is only slightly undulating. The collar is
cylindrical, with a length of 0.35 total length, and with 6
rectangular windows with rounded edges. The beams and
mullions are exceptionally heavy. The bowl is relatively
squat, acorn-shaped, rounding from the neck to 1.08 oral
diameters at 0.47 total length from the rim, then contracting
(40 in the upper half, 85 below, and near the aboral end
125 ). The aboral end is blunted.

The bowl has a major equatorial row of 8 large, squarish
to oval fenestrae, and a double row of minor ones; the upper,
minor ones form a row of approximately 24 elongate-oval

34

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

ones, and the distal row has about 10 somewhat larger,
rounded ones. The latter row is less regular than either of
the others. A few isolated, scattered lenestrae also occur
here and there. The interfenestral surface is reticulated, and
the framework of the collar has large primary alveoles. The
collar windows are all open.

Length, 62^.

Loricae of Dictyocysta lepida are commonly confused
with those of the more frequent D. reticulata, which species
has a single row of fenestrae, a flatter-sided middle bowl,
and a relatively longer collar. Dictyocysta lepida is not so
squat as D. lata, so big as D. grandis, nor so pointed aborally
as D. mexicana.

Recorded from two stations in the Pacific, as follows: one
(109) in the North Pacific trade region, and one (137) in
the California region.

There are 1 pump and 1 net sample, the former taken at
100 meters, the latter at 50 meters. Frequency, minimum.

Temperature: pump sample ig?8i, net sample 24? 84.
Salinity: pump sample 34.86, net sample 35.12. Density:
pump sample 24.72, net sample 23.50. pH: pump sample
8.18, net sample 8.34.

Dictyocysta magna Kofoid and Campbell
Dictyocysta magna Kofoid and Campbell, 1929, p. 294, fig. 569.

The tall, wide lorica, with elegans-Yike fenestration and
conical bowl, has a length of 1.44 oral diameters. The oral
margin is only slightly undulating and has a thin vertical
list. The collar has a length of 0.48 total length and is sub-
cylindrical, the diameter at the throat being 1.12 oral diam-
eters. It has an upper row of 8 squarish windows with
rounded edges, and a lower row of 10 subrectangular ones
which are subequal to those of the upper row. The beams
and mullions are hyaline, rather heavy, rounded, and
thick. All are open. The bowl is acorn-like with a trace of
shoulder, the upper seven-tenths being within 52 , and
the remaining part 113 . The aboral end is obtuse and
blunted.

The wall of the bowl has an equatorial row of 12 large,
subequal, subcircular fenestrae, and a second row of smaller
ones in similar number below the major row. A third row
of 6 yet smaller ones is below the second row, and a cluster
of tiny ones is found near the aboral end. The interfenestral
space is filled with tiny, subequal, prevailingly subcircular
areas. All fenestrae are closed by thin, hyaline panes.

Length, 75 to 95U.

Dictyocysta magna is unusually large. It differs from D.
mulleri not only in general surface pattern, but also in the
shape of the collar. It has a more pointed bowl than either
D. ampla or D. inaequalis, and is unlike all three species in
its collar fenestrae, all of which are subequal. It is not so
large as D. grandis, but has a double row of collar fenestrae
and not a single row as in grandis.

Recorded from two stations (3, 5-6) in the Atlantic drift.

There is 1 net sample from the surface, and 1 pump
sample from 50 meters. Frequency, 9 per cent at station 3.

Temperature (station 3 only), 14^66; salinity, 35.96; den-
sity, 26.81; pH, 8.19.

Dictyocysta minor Jorgensen
Dictyocysta minor, Kofoid and Campbell, 1929, p. 295, fig. 550.

The small, stout lorica, with mitrahke fenestration but
with small subuniform windows near the bowl, has a length
of 1.39 oral diameters. The oral margin has 8 flat arches.
The collar is truncated subcorneal, flaring (15°), straight-
sided, with a diameter of 0.92 oral diameter at the neck, and
with 8 subequal, rounded rectangular windows. The bowl
is plump, reaching 1.08 oral diameters at or near the middle.
The aboral end is obtusely pointed.

The bowl has 4 vertical, unequal rows of subpentagonal,
subhexagonal, or even squarish fenestrae, 10 to 12, 8 to 10,
12 to 14, 6 to 9, respectively, in number horizontally. In-
terpolated are minute fenestrae, and near the aboral end are
clustered about 40 very tiny ones. The framework of the
collar is heavy, thick, and lifted above the surface. The
windows have neat frames, and appear to be closed with
hyaline panes.

Length, 47 to 54^1.

Dictyocysta minor, though it has the mitra type of wall
structure, differs from D. mitra not only in having aboral
clusters of small fenestrae, but also in being shorter and
wider. Dictyocysta dilatata is somewhat similar but has an
aboral spine, more flaring collar, and slimmer proportions,
and lacks the small fenestrae.

Recorded from sixteen stations, ten in the Atlantic and six
in the Pacific, as follows: one (3) in the Atlantic drift, three
(14, 15, 16) in the Gulf Stream, three (18, 19, 20) in the
Sargasso Sea, three (22, 23, 24) in the Atlantic equatorial
region, two (46, 47) in the Galapagos region, three (63, 64,
65) in the South Pacific middle latitudes, and one (no) in
the North Pacific trade region.

There are 9 pump and 1 1 net samples, of which 2 were
taken at the surface, 5 at 50 meters, and 13 at 100 meters.
Maximum frequency, 10 per cent at station no; other records
above minimum (all 3 per cent) from stations 16, 18, 63, 64;
average in Atlantic net samples, 4.2 per cent.

Temperature: Atlantic, pump samples i4?02-23?64( 1 8?35),
net samples i4?02-24?io (19? 19); Pacific, i5?03-i7?96
(i6?i8) and 23?30-23?88 (23^59), respectively. Salinity:
Atlantic, pump samples 35.10-36.48 (36.00), net samples
35.59-37.05 (36.29); Pacific, 34.30-34-73 (34-53) an d 35-32-
35.96 (35.64), respectively. Density: Atlantic, pump samples
24.84-26.66 (25.97), net samples 24.49-26.81 (25.76); Pacific,
25.10-25.48 (25.46) and 24.11-24.42 (24.36), respectively.
pH: Atlantic, pump samples 8.06-8.23 (8.16), net samples
8.06-8.25 (8.17); Pacific, 8.16-8.23 (8.19) and 8.08-8.14
(8.10), respectively.

Dictyocysta mitra Haeckel emended Kofoid and Campbell

(Figures 60, 61)
Dictyocysta mitra, Kofoid and Campbell, 1929, p. 296, fig. 548;
Hofker, 1931, pp. 381-382, figs. 74-75.

The relatively plump lorica, with scant division between
collar and bowl, large, window-like meshwork, and rounded
aboral end, has a length of 1.47 oral diameters. The oral
rim is arched with 7 (8) broad frames. The collar region is
generally biconical (33 in the upper and 35° in the lower

TAXONOMY AND DISTRIBUTION

35

half); its least diameter, 0.88 oral diameter, is at the junction
of the upper and lower sections. At the neck the diameter
equals that of the oral rim. The bowl expands from the
throat to a diameter of 1.13 oral diameters at 0.54 total
length from the rim, the collar being 0.33 total length. The
lower bowl rounds off below the maximum diameter, and
the aboral end is blunt to rotund.

The wall is extremely thin. The whole surface is fenes-
trated, with the upper windows squarish. The fenestrae of
the second row are smaller and more irregular, and the
fenestrae of the bowl are less uniform, 5 to 7 in vertical rows
and 10 to !2 in each horizontal row; at the aboral end is a
nest of tiny ones, and odd minute ones are scattered among
the larger ones of the bowl. All fenestrae are enclosed by
thickened beams and mullions, with a fine molding around
each window. All windows are closed by transparent panes.

Length, 60 to 8ou.

Hofker's ( 1 93 1 ) material appears typical although his
figures are generalized.

Dictyocysta mitra is less stout and has a narrower aboral
end than D. obtusa. It has a less distinctly set-off and less
flaring collar region than D. dilatata (some of the Carnegie
loricae, as fig. 61, approach dilatata in this character), and
less acute aboral end. The fenestration differs markedly
from that of D. jenestrata, as does the general shape of the
bowl; in jenestrata the bowl is much less elongate. Dictyo-
cysta californiensis is smaller and has a greatly swollen bowl;
D. spinosa and D. extensa have oral spines lacking in mitra.

Recorded from twenty-one stations, six in the Atlantic
and fifteen in the Pacific, as follows: two (2, 15) in the Gulf
Stream, three (3, 4, 5) in the Atlantic drift, one (23) in the
Atlantic equatorial region, four (42, 43, 44, 68) in the Gala-
pagos region, four (61, 63, 64, 65) in the South Pacific
middle latitudes, three (in, 128, 144) in the North Pacific
middle latitudes, and four (131, 133, 134, 135) in the Cali-
fornia region.

There are 18 pump and 11 net samples, of which 1 was
taken at the surface, 9 at 50 meters, 18 at 100 meters, and 1
at 1000 meters. Maximum frequency, 6 per cent at station
65; other records above minimum (2 to 3 per cent) from
stations 3, 4, 44, 63, 64, in, 131; averages, 2.1 and 2.5 per
cent in Atlantic and Pacific net samples, respectively.

Temperature: Atlantic, pump sample i8?56, net samples
i3? 7 9-20?35 (i5?o6); Pacific, 3 ? 9 8-i 5 ?o3 (n?36) and
io?23-i9?97 (i5?4o), respectively. Salinity: Atlantic, pump
sample 36.45, net samples 35.88-36.41 (36.02); Pacific, 33.07-
35.04 (34.46) and 33.36-35.04 (34.25), respectively. Density:
Atlantic, pump sample 26.26, net samples 25.76-27.01 (26.72);
Pacific, 24.64-26.19 (25.25) and 25.31-27.25 (26.04), respec-
tively. pH: Atlantic, pump sample 8.19, net samples 8.10-
8.21 (8.15); Pacific, 7.88-8.39 (8.15) and 7.76-8.32 (8.02),
respectively.

Dictyocysta mulleri (Imhoff) Jorgensen emended Kofoid

and Campbell
Dictyocysta midleri, Kofoid and Campbell, 1929, pp. 296-297,

fig- 572-
The short lorica, with spool-shaped collar with double

row of windows, and angled, aborally pointed bowl, has a
length of 1.51 oral diameters. The oral rim is undulating,
with 6 very fiat arches. The collar is spool-like, contracting
from the rim to 0.94 oral diameter near 0.48 oral diameter
from the rim, and then expanding to Ml oral diameters at
the neck, which is 0.81 oral diameter below the rim. There
is a double row of open windows, the upper row being ol 6
squarish ones, the lower being formed by 9 rounded rec-
tangles. The beams and mullions are thick and heavy-set.
The bowl is subcorneal (55 in the upper two-thirds, in-
creasing to 115° in the lower section). There is a rounded
but short shoulder below the collar, the maximum diameter
of which is 1.16 oral diameters. The aboral end is obtuse
and blunted.

The wall of the bowl has an anterior row of 24 double,
circular fenestrae, and this row is followed by an incomplete
row of smaller circles. A row of 10 to 12 large, oval, post-
equatorial fenestrae adorns the bowl, and a lower row of 4
to 6 small windows is also present. The remaining surface
is covered with much smaller, relatively thick-walled fenes-
trae, although the area near the large fenestrae is free of tiny
circles. The fenestrae all have clear panes.
Length, 55 to 631.1.

Dictyocysta mulleri differs from all other species in the
shape of the aboral end, which, combined with its wall char-
acters, differentiates it at once with little difficulty. Its
nearest relative is, possibly, D. magna, which not only is
longer, but also lacks the spool-like collar. Dictyocysta
ampla has a less angled bowl.

Recorded from three stations, two in the Atlantic and one
in the Pacific, as follows: one (16) in the Gulf Stream, one
(20) in the Sargasso Sea, and one (82) in the region of
South Pacific island fields.

There are 2 pump and 1 net sample, all taken at 100
meters. Frequency, minimum.

Temperature: 19^62, 26^02, and 24?34 in the Atlantic
pump and net samples and Pacific pump sample, respectively.
Salinity: 36.48, 36.51, and 36.46 in the Atlantic pump and
net samples and Pacific pump sample, respectively. Density:
26.01, 24.19, and 24.65 in the Atlantic pump and net samples
and Pacific pump sample, respectively. pH: 8.17, 8.32, and
8.19 in the Atlantic pump and net samples and Pacific pump
sample, respectively.

Dictyocysta nidulus Kofoid and Campbell
Dictyocysta nidulus Kofoid and Campbell, 1929, pp. 297-298,

fig. 565.
The fairly tall lorica, with tiara-like fenestration and a
long bowl, has a length of 1.6 oral diameters. The oral
margin is simple and has a narrow list. The collar is 0.4
total length in length, virtually a cylinder, with a single row
of 7 rectangular (width 0.53 height) windows; in some indi-
viduals the windows, which are always open, have single,
somewhat oblique crossbars which subdivide them into two
more or less equal sections. The beams and mullions are
thin, rounded, and hyaline. The bowl is acorn-like in shape,
with some swelling a little below the collar (reaching 1. 1
oral diameters in diameter), and below it becomes angular

36

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

subcorneal (42 in the flat-sided upper seven-tenths and 114
in the lowermost rounded section). The aboral end is ob-
tusely rounded.

The wall of the bowl has an irregular row of 10 closed,
subrectangular, large fenestrae and a pre-equatorial row of
large Syracosphaera. Minute circular areas (coccoliths? )
cover the remainder of the surface.
Length, 75U.

Dictyocysta nidulus resembles D. reticulata but diflers
mainly in proportions, regularity of large fenestrae, presence
of large Coccolithophoridae, and general shape of bowl.
Dictyocysta mexicana has greater angularity, more projecting
aboral end, more localized fenestrae, and shorter length (57
to 66u).

Recorded from one station (4) in the Atlantic drift, in a
net sample taken at 50 meters. Frequency, 6 per cent.

Temperature, 14^32; salinity, 36.00; density, 26.91; pH,
8.15.

Dictyocysta occidentalis Kofoid and Campbell
Dictyocysta occidentalis Kofoid and Campbell, 1929, pp. 298-

299, fig. 556.
The fairly small, symmetrical lorica, with mitra-Uke fenes-
tration, lepida-\\kt collar, and globose bowl, has a length of
1.57 oral diameters. The oral margin is undulating, with 7
arches. The collar is subcylindrical, 0.63 of the length of the
bowl in length, with 7 subquadrangular, closed windows,
and thick beams and mullions. The bowl is globular, its
greatest diameter, at the middle, being 1.15 oral diameters.
There is a rounded shoulder just below the throat. The
aboral end is hemispherical.

The wall of the bowl has an equatorial zone of 8 to 10
subequal, round fenestrae, a post-equatorial one of 6 to 8,
and an aboral group of very small ones. The meshwork is
heavy and pronouncedly minutely alveolar. Coccoliths are
often found, especially in the aboral fenestrae.
Length, 55 to 62^1.

Dictyocysta occidentalis has a globose bowl somewhat like
that of D. polygonata, but the latter species has large circles
forming the wall of the bowl, and has no equatorial or other
limited zones of fenestrae. Dictyocysta duplex has "duplex"
walls and a very different collar. Dictyocysta tiara has
thinner beams and an ovoid bowl.

Recorded from three stations in the Pacific, as follows:
one (136) in the California region and two (141, 144) in
the North Pacific middle latitudes.

There are 3 pump samples, of which 1 was taken at the
surface and 2 at 50 meters. Frequency, minimum.

Temperature, i8?6cr-25?3i (22?86); salinity, 34.64-35.38
(35.09); density, 23.46-24.84 (24.02); pH, 8.33-8.37 (8.34).

Dictyocysta pacifica Kofoid and Campbell
Dictyocysta pacifica Kofoid and Campbell, 1929, p. 299, fig. 555.
The short lorica, with the general shape of mitra as well
as with similar fenestration, but with oral denticles, has a
length of 1.43 oral diameters. The oral margin is regularly
set with about 40 sparsely spaced, triangular denticles. The
collar is a segment of an inverted cone (io°) with a length

of 0.57 total length, a diameter of 0.86 oral diameter at the
throat, and 6 squarish windows, each set into a relatively
wide frame. The windows are all closed, and the beams
and mullions are rather thick. The collar merges with the
plump bowl. The bowl expands from the throat to 1.14
oral diameters near its equator. Below this it is convex
conical (6o° in the anterior part, increasing to 125 in the
lower half). The aboral end is pointed.

The wall of the bowl has 2 rather regular and 1 irregular
row of fenestrae. The fenestrae of the first row, 10 in num-
ber, are similar in size to those of the second row, which
number 12; both sets are subcircular in shape and subequal
in size. The fenestrae of the distal set form an irregular
row of 6 small windows. The interfenestral area has smaller
polygons; coccoliths are not rare in this distal region.
Length, 41 to 52U..

Dictyocysta pacifica differs from the species of the mitra
type in having oral denticles; D. extensa and D. spinosa,
however, have larger spinelike teeth instead of tiny spikes.
In some ways D. pacifica recalls D. minor, but it has smaller
fenestrae in the distal group, as well as oral denticles.

Recorded from four stations in the Pacific, as follows: two
(35, 35-36) in the Pacific equatorial region, one (45) in the
Galapagos region, and one (136) in the California region.
There are 3 net samples and 1 pump sample, of which 1
was taken at the surface, 1 at 50 meters, and 2 at 100 meters.
Frequency, 2 per cent at station 136; other records minimum.
Temperature: pump sample i8?8y, net samples i6?30-
21^69 (i8?99). Salinity: pump sample 35.02, net samples
34.88-35.21 (35.04). Density: pump sample 25.09, net
samples 24.48-25.60 (25.04). pH: pump sample 8.39, net
samples 7.92-8.12 (8.02).

Dictyocysta reticulata Kofoid and Campbell

Dictyocysta reticulata Kofoid and Campbell, 1929, p. 300, fig.

560; Marshall, 1931, pp. 653-654.
Dictyocysta lepida, Hofker, 1931, pp. 382-384, figs. 76-78.

The plump lorica, with arched collar windows in a single
row resembling the columns of a Greek temple, wide, low
bowl, rounded aboral end, and large oval fenestrae in the
bowl, has a length of 1.37 oral diameters. The oral margin
has 6 (7) flattened arches. The collar is nearly cylindrical.
0.46 total length in length, and there are 6 (7) tall, subequal
windows. The subhemispherical bowl reaches 1.15 oral
diameters at 0.5 total length, and thence more or less evenly
rounds to the unmodified aboral end.

The wall of the bowl is hardly 0.03 oral diameter in
thickness. The whole bowl is fenestrated with subuniform,
rounded, deeply pitted polygons except for a single row of
large subequal, closed, oval fenestrae, 6 (7) in number, which
are in the bowl's equatorial region. The beams and mullions
of the closed fenestrae of the collar are hyaline, rounded, and
squarish in form.

Length, 55 to 65(1.

Hofker's material probably includes several related species,
as this writer does not closely differentiate the species of this
group.

TAXONOMY AND DISTRIBUTION

37

Dictyocysta reticulata is shorter and has a longer bowl
than D. grandis; it has no aboral point like D. mexicana;
and its howl is more angled at the shoulder than is that of
D. lata. Most of the remaining species either are of the D.
mitra type, or else have 2 rows of clear-cut windows in the
collar. This species is by far the most common in warm
water.

Recorded from forty-seven stations, seventeen in the
Atlantic and thirty in the Pacific, as follows: three (2, 14.
[6) in the Gulf Stream, three (3, 4, 5) in the Atlantic drift,
two ( t8, 19) in the Sargasso Sea, seven (22, 23, 24, 25, 26,
28, 30) in the Atlantic equatorial region, two (33, 34) in the
Caribbean Sea, five (35, 35-36, 36, 37, 154) in the Pacific
equatorial region, tweb'e (40, 41, 42, 44, 45, 47, 69, 71, 77,
78, 79, 80) in the Galapagos region, three (48, 84, 92) in the
region of South Pacific island fields, five (59, 62, 64, 65, 67)
in the South Pacific middle latitudes, two (109, no) in the
North Pacific trade region, one (in) in the North Pacific
middle latitudes, and two (133, 149) in the California region.
Dictyocysta reticulata is apparently absent only in cool
regions such as the North Sea, the American cold-water
region, the East Asiatic marginal sea, and the Alaskan
secondary region.

There are 34 pump and 44 net samples, of which 19 were
taken at the surface, 31 at 50 meters, and 28 at 100 meters.
Maximum frequency, 13 per cent at station 77; other records
above minimum (2 to 11 per cent) from stations 2, 3, 22, 24,
25, 26, 28, 35, 35-36, 40, 41, 42, 44, 45, 47, 48, 59, 62, 65, 69,
78, 80, 154; averages, 1.0 and 2.1 loricae in Atlantic and
Pacific pump samples, and 2.8 and 4.1 per cent in Atlantic
and Pacific net samples, respectively.

Temperature: Atlantic, pumpsamples i4?95~28?25 (23?46),
net samples i3?37~27?88 (20? 12); Pacific, i3?93~27?99
(2^23) and i4?33-27?52 (2^05), respectively. Salinity:
Atlantic, pump samples 29.70-37.15 (35.33), net samples
35.61-36.81 (36.15); Pacific, 31.62-36.17 (34.86) and 31.62-
36.44 (34.48), respectively. Density: Atlantic, pump samples
18.62-26.08 (23.89), net samples 23.26-27.01 (25.94); Pacific,
20.20-26.21 (24.37) ano - 20.34-26.17 (24.47), respectively.
pH: Atlantic, pump samples 8.14-8.31 (8.21), net samples
7.93-8.30 (8.16); Pacific, 7.85-8.37 (8.17) and 7.87-8.38
(8. n), respectively.

Dictyocysta speciosa Jorgensen
Dictyocysta speciosa, Kofoid and Campbell, 1929, pp. 300-301,
fig- 575-

The large lorica, with elegans-like fenestration and aborally
constricted bowl, has a length of 1.63 oral diameters. The
oral margin is mildly arched, with 8 flattened rims. The
collar (1.0 oral diameter in length) is subcylindrical, the
diameter at the lower end being 1.12 oral diameters. The
collar has a double row of fenestrae. The upper row is made
up of 8 rectangular windows with squarish corners, larger
than the 9 subpentagonal to square ones of the lower row.
The beams and mullions are hyaline and rounded, and en-
close the open windows. The bowl has a considerably
rounded shoulder, reaches 1.31 oral diameters a little below

the lower end of the collar, then rapidly contracts (53°). and
becomes almost hemispherical (125 ) within its lower fourth.
The upper section is flat-sided. The aboral end is obtusely
blunted.

The wall of the bowl has a postequatorial row oi 8
rather small round fenestrae, and a lower row ot 6 even
smaller oval ones. The interfencstral area has many small
circles of nearly uniform size.

Length, 66 to 8op.

Dictyocysta speciosa is larger than other species related to
D. elegans, and has a bowl shaped unlike those in the re-
mainder of that general group. Once seen, it is not hard
to recognize.

Recorded from five stations (3, 4, 5-6, 6, 6-7) in the
Atlantic drift and the North Sea.

Net samples only, of which 8 were taken at the surface, 2
at 50 meters, and 4 at 100 meters. Maximum frequency, 40
per cent between stations 5 and 6; all other records above
minimum (2 to 36 per cent), except between stations 6 and
7, where only 1 lorica was found.

Temperature, ii?28-i4?32 (i2?8o); salinity, 35.51-36.06
(35.77); density, 26.91-27.15 (27.00); pH, 7.90-8.15 (8.10).

Dictyocysta spinosa Kofoid and Campbell
Dictyocysta spinosa Kofoid and Campbell, 1929, p. 301, fig. 553.

The rather stout lorica, with mitra-Yike fenestration and
distinct oral spines, has a length of 1.36 oral diameters. The
oral rim has 8 flattened arches; from the middle of each
arch arises an outspread needle-like spine. The collar is in-
verted subconical and flaring (8°); its length is 0.3 total
length. The diameter at the throat equals that of the rim,
and the sides are concave. There are 8 rounded, rectangular,
closed windows. The bowl is broadly ovate, has no shoul-
der, and reaches 1.24 oral diameters near its middle. The
aboral end is generally blunted.

There are 8 rows of more or less irregular rows of uneven
fenestrae in the bowl. These increase in number but de-
crease in size toward the middle.

Length, 42 to 46(1.

Dictyocysta spinosa and D. extensa are the only species
with oral spines, and the latter species differs from spinosa
in the altogether different wall structure, having fewer large
fenestrae, and many small polygons on the longer, more
acute bowl.

Recorded from four stations, two each in the Atlantic and
the Pacific, as follows: one (19) in the Sargasso Sea, one
(23) in the Atlantic equatorial region, one (75) in the Gala-
pagos region, and one (135) in the California region.

There are 1 pump sample and 3 net samples, all taken at
100 meters. Frequency, minimum.

Temperature: Atlantic, net sample 22?42; Pacific, pump
sample i8?95, net sample i8?4o. Salinity: Atlantic, net
samples 34.88-37.05 (35.85); Pacific, pump sample 34.88,
net sample 35.47. Density: Atlantic, net sample 25.67;
Pacific, pump sample 24.96, net sample 25.55. pH ; Atlantic,
net samples 8.18-8.25 (8.21); Pacific, pump sample 8.34, net
sample 8.10.

38

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Dictyocysta tiara Hacckel
Dictyocysta tiara, Kofoid and Campbell, 1929, p. 302, fig. 567.

The rather moderately tall, narrow lorica, with single row
of fenestrae and elongated bowl, has a length of 2.0 oral
diameters. The oral margin is undulating, with 8 very flat
arches. The collar is cylindrical, 0.75 oral diameter in length,
and has 8 rectangular, subequal open windows with rounded
lower and angular upper corners. The bowl has a narrow
shoulder, reaches a diameter of 1.25 oral diameters at the
shoulder, which is a little lower than the lower end of the
collar, then contracts (30 ) for two-thirds of its length, and
finally becomes subcorneal (98 ). The aboral end is rounded.

The wall of the bowl has a single row of 8 elongated,
narrow oval fenestrae in the equatorial region. More
numerous tiny fenestrae occur in a double row below the
major row. The remaining surface of the bowl and the
supporting beams and mullions of the collar are studded
with closely set, minute coccoliths. The collar windows are
open.

Length, 8o|i.

This much misunderstood species is more elongated and
narrower than any other.

Recorded from four stations in the Pacific, as follows: one
(45) in the Galapagos region, one (64) in the South Pacific
middle latitudes, and two (in, 112) in the North Pacific
middle latitudes.

There are 5 pump samples, of which 1 was taken at the
surface and 2 each at 50 and 100 meters. Frequency, 2 per
cent at stations 45, 64, m; average, 1.6 per cent.

Temperature, i5?oo-22?43 ( ig?gi ); salinity, 34.54-35.26
(34.73); density, 24.02-25.43 (24.58); pH, 8.10-8.22 (8.16).

COXLIELLIDAE Kofoid and Campbell emended
Coxliellidae, Kofoid and Campbell, 1929, p. 91.

The family includes four genera: Coxliella, Climacocylis,
Helicostomella, and Metacylis. Only the last does not occur
in the material of this expedition. All but Helicostomella
are eupelagic, the latter being coastal; most of them are
tropical.

Coxliellinae Kofoid and Campbell
Coxliellinae Kofoid and Campbell, 1939, p. 90.

The Coxliellinae include those members of the family in
which the spiral lamina extends the whole length of the
lorica, except in some species in which it may be limited.

The genera included are Coxliella and Climacocylis.

COXLIELLA Brandt emended
Coxliella, Kofoid and Campbell, 1929, pp. 95-96.

Coxliella has a spiral lamina which forms the bowl and
horn. This spiraling is also found in the collar of Codonel-
lopsis and in a few species of Tintinnopsis. It is not un-
likely that Coxliella had its origin in the latter genus, and
that the spiral collar of Codonellopsis is of a different source.
The wall is rigid, the flaccidity of that of Climacocylis being
derived. The aboral end of Coxliella is closed, that of

Climacocylis open; this is an additional suggestion of the
derivation of the latter genus.

Species of Coxliella are common in warm and temperate
seas. Only the subgenus Protocochliella is from cool waters
in both hemispheres, and is not found in the tropical ocean.

Six species are described here.

Coxliella declivis Kofoid and Campbell
Coxliella declivis Kofoid and Campbell, 1929, p. 97, fig. 192.

The tapering subconical lorica, with few turns and narrow
aboral horn, has a length of 1.89 to 1.93 oral diameters. The
oral margin is entire, thin, and smooth. The short, stout
bowl is cylindrical in the upper fourth, and convex conical
in the lower three -fourths; the upper section is 55 to 6o°,
whereas near the horn it is 95 . The short, conical aboral
horn is somewhat under 0.3 oral diameter in length; it is
twisted, and its end is minutely blunted.

The wall reaches as much as 0.09 oral diameter in thick-
ness across the widest of the 9 to 10 left-turning spiral whorls.
The fifth and sixth turns reach a width of 0.33 oral diameter;
the uppermost and lowermost ones are only half that much,
or less. There are thin laminae with three to five layers of
tiny alveoles closely packed within their confines. The wall
is dull, and lacunae are absent.

Length, nou,.

Coxliella declivis is more tapering and has a more sym-
metrical aboral horn than C. laciniosa. Its wide turns sug-
gest C. pseudannulata, but its horn is decidedly different,
and neither is it like that of C. pelagica. Coxliella decipiens
is longer, with more and narrower spiral turns and some-
what similar aboral end. The remaining species are all so
different that no difficulty in identification should arise.

Recorded from seven stations, two in the Atlantic and
five in the Pacific, as follows: two (18, 19) in the Sargasso
Sea, two (41, 85) in the Galapagos region, one (99) in the
Pacific equatorial region, and two (133, 134) in the Cali-
fornia region.

There are 2 pump and 6 net samples, of which 2 were
taken at the surface, 2 at 50 meters, and 4 at 100 meters.
Maximum frequency, 3 per cent at station 99; other records
at 2 per cent from stations 19, 85.

Temperature: Atlantic, net samples 20?32-25?3i (22?68);
Pacific, pump samples i8?i8-22?68 (20^43), net samples
20?42-27?89 (25^38). Salinity: Atlantic, net samples 36.81-
37.15 (37.00); Pacific, pump samples 34.63-34.70 (34.66),
net samples 34.19-36.24 (35.15). Density: Atlantic, net
samples 24.80-26.07 (25.48); Pacific, pump samples 23.82-
24.96 (24.39), net samples 22.50-24.06 (23.31). pH: At-
lantic, net samples 8.21-8.27 (8-24); Pacific, pump samples
8.34-8.47 (8.40), net samples 8.1 1-8.22 (8.18).

Coxliella fasciata (Kofoid) Brandt
(Figures 30, 34)

Coxliella fasciata, Kofoid and Campbell, 1929, pp. 97-98, fig.

205.

The greatly elongated conical lorica, with few spiral turns
and slight flare, and with narrowed aboral end, has a length

TAXONOMY AND DISTRIBUTION

39

of 3.62 oral diameters. The oral margin is thin-edged, en-
tire, and rolled over. The upper end of the howl flares as a
cone (6o°) and includes the uppermost of the spiral turns.
Its sides are outwardly concave. The greater part of the
remaining portion of the bowl is also conical (io°), except
the aboral fifth, which contracts (16°) somewhat more.
The aboral end is not prolonged or especially modified as a
horn, save as it twists a bit.

The wall is made up of 10 left-wound spiral turns. The
uppermost 5 are 0.25, 0.44, 0.53, 0.44, and 0.37 oral diam-
eter, respectively, in width. The aboral 5 are more or less
subuniformly about 0.44 oral diameter. Optically dark lines
separate the turns, and they rest one against another without
overlap. The wall is relatively thick, being 0.09 oral diam-
eter in the thickest turn. There are thin laminae which
enclose three to seven layers of rather tiny ovoid alveoles.
The wall is almost transparent. The cavity follows the outer
contour to the very tip; sometimes the aboral end is minutely
open.

Length, 3i2(i.

A peculiarly shaped, perhaps abnormal lorica is figured
(fig. 30) in addition to the more usual type (fig. 34). This
much shorter lorica has 19 almost horizontal turns, and a
wavy, eccentric aboral third. It may be a young individual
which for some reason did not finish out its lorica, or may
be a representative of a rarer species not yet reported.

Coxliella fasciata is easily differentiated from the other
species of the genus because of its elongated conical form
with few turns. Its aboral end is pointed, unlike that of
C. minor, which species is not tropical as is -fasciata. Its con-
tour and aboral end differ from those of C. helix; it is not
so stout as C. decipiens, and lacks the solid horn which that
species has.

Recorded from six stations, five in the Atlantic and one
in the Pacific, as follows: one (14) in the Gulf Stream, one
(20) in the Sargasso Sea, two (23, 24) in the Atlantic equa-
torial region, one (34) in the Caribbean Sea, and one (139)
in the North Pacific trade region.

There are 1 pump and 5 net samples, of which 1 was
taken at the surface, 2 at 50 meters, and 3 at 100 meters.
The pump sample was taken in the Pacific and the net
samples in the Atlantic. Frequency, 4 per cent at station 14;
other records all minimum; average, 2 per cent.

Temperature: pump sample 26^72, net samples 14^02-
24?98 (i9?62). Salinity: pump sample 34.82, net samples
35.59—36.73 (36.10). Density: pump sample 22.69, net
samples 24.52-26.66 (25.45). P H: pu m P sample 8.34, net
samples 7.96-8.21 (8. 11).

Coxliella helix (Claparede and Lachmann) Brandt emended
Kofoid and Campbell

Coxliella helix, Kofoid and Campbell, 1929, pp. 99-100, fig.
204.

The tall, tubular, vaselike lorica, with toothed oral margin,
regular contour, and short, conical aboral horn, has a length
of 3.72 oral diameters. The oral margin is regularly serrate,
with about 36 short, evenly spaced, triangular, outspread

teeth. The long howl contracts below the margin as a basal
segment of an inverted concave cone (45°) with a width of
about 0.1 total length. The tubelike part of the bowl oc-
cupies somewhat less than 0.6 total length; it is without local
bulges or contractions. The posterior part is inverted, de-
cidedly convex conical (22 in the upper two-thirds and <><■
in the lower third). The conical (22' ) aboral horn has a
length of only 0.5 oral diameter.

The spiral lamina which forms the wall has 18 left-turning
spirals. These bands are widest across the mid-region of the
bowl and subuniformly narrower in the upper and lower
parts. At their widest they reach nearly 0.3 oral diameter.
They tend to rest one on another in such fashion that the
lateral contour presents a saw-toothed edge. An optically
dark line marks their boundaries. The wall is subuniformly
0.02 oral diameter in thickness except near the suboral flare,
where it gradually thins to the serrate margin. The wall has
distinct, large, hexagonal secondary structures with enclosed
minute alveoles. Toward the aboral end, adherent irregular
blobs occur. The aboral horn is hollow and the lumen
agrees with the outer contour exactly.

Length, 19611.

The Carnegie lorica is in so many ways different from
that figured by Kofoid and Campbell that it might, with
some justification, have been assigned to a new species. The
regularity of fades, presence of teeth, length of bowl, and
shortness of horn are all characters of note. Yet the two
loricae seem to be essentially similar and to have definable
characters in common.

Coxliella helix bears little resemblance to most of the
species, being taller and more slender. It is somewhat like
C. fasciata, but has a tubular bowl and definite contracted
aboral horn, as well as suboral flare, and a toothed margin.
Coxliella minor is also elongated, but its aboral horn is bul-
bous. Coxliella decipiens is relatively wider, and of alto-
gether different appearance. Coxliella helix is not likely to
be confused with others.

Recorded from three stations in the Atlantic, as follows:
two (19, 20) in the Sargasso Sea and one (23) in the Atlantic
equatorial region.

There are 3 net samples, of which 2 were taken at 50
meters and 1 at 100 meters. Frequency, 3 per cent at station
19.

Temperature, 2o?99-25?3i (22?95); salinity, 36.04-37.15
(36.64); density, 24.89-25.38 (25.28); pH, 8.14-8.27 (8.20).

Coxliella Iaciniosa (Brandt) Brandt emended Kofoid

and Campbell

(Figures 29, 31, 32, 33)

Coxliella Iaciniosa, Kofoid and Campbell, 1929, p. 100, fig. 193;
Marshall, 1934, p. 642.

The generally short, stout, conical lorica, with entire rim
and short, twisted, conical aboral horn, has a length of 1.21
oral diameters. The oral margin is thin-edged, erect, and
smooth. The stout, generally conical bowl is subcylindrical
in the anterior 0.59, and convex conical aborally (at first
32 , and 105° nearer the horn). The sides are full and the

4°

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

spiral turns bulge somewhat. The aboral horn is short (0.17
oral diameter), conical (33°), and twisted.

The wall is thicker suborally than in the lower bowl and
hollow horn; the thickness is 0.05 oral diameter at the maxi-
mum, and less than half as much elsewhere. Each of the
spiral turns is enclosed by a thin inner and outer lamina,
and these, in turn, enclose two to five layers of rounded
alveoles. The left-turning spiral turns number 7 to 10. They
usually are more or less subuniformly 0.21 oral diameter in
width, save for the fourth and fifth ones, which reach 0.26.
Optically dark lines separate the turns, and oval lacunae in
1 or 2 rows occur near the aboral end of the widest turn;
sometimes a few odd ones are found on the turn just above
this one. Very minute, or primary, alveoles may be found
under the highest magnification of the microscope, but in
general the wall is pallid.

Length, 75 to 140^1.

Marshall (1934) finds loricae with but a single layer of
alveoles, and with aboral swelling. The Carnegie loricae
differ among themselves considerably, and a few extremes
are figured (figs. 31, 32, 33); figure 29 shows a more com-
mon form. One of these specimens (fig. 32) has some like-
ness to Coxliella declivis, but differs in important respects.

Coxliella laciniosa is most like C. declivis, but its walls are
different, the spiral turns are unlike, and the aboral horn is
not blunted. Coxliella deapiens is longer and relatively nar-
rower, with more turns and a different horn. The remain-
ing species, in spite of the great variability of individual
loricae of laciniosa, are quite unlike it and never lead to
confusion.

Recorded from thirty-seven stations, twelve in the Atlantic
and twenty-five in the Pacific, as follows: two (15, 16) in the
Gulf Stream, four (17, 18, 19, 20) in the Sargasso Sea, five
(23, 25, 27, 29, 30) in the Atlantic equatorial region, one
(34) in the Caribbean Sea, six (43, 45, 46, 47, 71, 78) in the
Galapagos region, six (48, 82, 84, 85, 95, 158) in the region
of South Pacific island fields, six (100, 102, 105, iog, 140,
150) in the North Pacific trade region, three (136, 146, 148)
in the Calitornia region, three (141, 144, 145) in the North
Pacific middle latitudes, and one ( 155) in the Pacific equa-
torial region.

There are 19 pump and 24 net samples, of which 4 were
taken at the surface, 22 at 50 meters, and 17 at 100 meters.
Maximum frequency, 10 per cent at station 145; other
records above minimum (2 to 4 per cent) from stations 15,
27, 34, 48, 84, 85, 145. 148, 155; average in Pacific net
samples, 2.4 per cent.

Temperature: Atlantic, pumpsamples 2^85-27? 11 (24?2o),
net samples i4?6o-25?54 (22?57); Pacific, i5?85-27?52
(21^63) and i8?87-28?74 (24^55), respectively. Salinity:
Atlantic, pump samples 36.22-36.60 (36.41), net samples
35-7°-37- r 5 (3 6 4 2 ); Pacific, 34.32-36.46 (35.23) and 34.32-
36.42 (35.08), respectively. Density: Atlantic, pump samples
23.62-25.49 (24.65), net samples 23.98-26.62 (25.10); Pacific,
22.71-25.60 (24.00) and 22.31-25.09 (23.63), respectively.
pH: Atlantic, pump samples 8.23-8.29 (8.26), net samples
7.93-8.30 (8.17); Pacific, 7.83-8.34 (8.19) and 8.13-8.39
(8.25), respectively.

Coxliella longa (Brandt) Laackmann
Coxliella longa, Kofoid and Campbell, 1929, p. 101, fig. 196.

The bullet-shaped lorica, with turns of moderate width,
rounded aboral end, and short, curved aboral point, has a
length of 2.1 oral diameters. The oral margin is irregularly
toothed with larger and smaller scattered denticles. The
bowl is subcylindrical in its upper three-fifths and becomes
convex conical in the lower part (35 ° in the anterior region
and 105 near the horn). At the aboral end is a slightly
curved, projecting point or horn; its free tip is minutely open.

The wall is only 0.03 oral diameter in thickness across the
widest turn. There are relatively thick radial prisms or
alveoles in two to four layers between the laminae. At the
free tip of the aboral end the wall is somewhat swollen
around the opening. Otherwise the cavity conforms to the
outer contour. Optically dark lines separate adjacent spiral
turns, of which there are 13. These become wider toward
the aboral end, the widest one being 0.3 oral diameter, which
is three times the width of the narrower anterior ones. Three
or four small oval lacunae, with their long axes directed up-
ward, are found on the widest turn.

Length, 130 to 135U.

Coxliella longa has more turns and a shorter aboral horn
than C. declivis, C. laciniosa, and C. pseudannulata. It also
has more turns than C. intermedia (an antarctic species),
and an aboral horn. Coxliella tubularis is bigger, with more
turns and a relatively better-developed horn.

Recorded from six stations in the Pacific, as follows: three
(47, 69, 78) in the Galapagos region, one (65) in the South
Pacific middle latitudes, and two (82, 83) in the region of
South Pacific island fields.

There are 1 pump and 5 net samples, of which 2 were
taken at the surface, 3 at 50 meters, and 1 at 100 meters.
Maximum frequency, 3 per cent at station 69; average, 1.5
per cent.

Temperature: pump sample 24^38, net samples 15^03-
27?46 (22?94). Salinity: pump sample 36.03, net samples
34.30-36.49 (35.86). Density: pump sample 24.33, net
samples 23.67-25.44 (24.18). pH: pump sample 8.14, net
samples 8.10-8.24 (8-17).

Coxliella pseudannulata (JSrgensen) Brandt
(Figure 28)

Coxliella pseudannulata, Kofoid and Campbell, 1929, p. 102,
fig. 191.

The moderately stout lorica, with wide, bulging median
spiral turns and thick, blunted aboral end, has a length of
2.6 oral diameters. The oral margin is minutely irregular
and thin-edged. The bowl is asymmetrically subcylindrical
in the anterior 0.45, conical (32 ) in the submedian 0.4, and
bluntly conical (26 ) in the lowermost section. The aboral
end is rounded off.

The wall is made up of 9 rather wide, right-wound spiral
turns, the width of the median 2 being 0.46 and 0.37 oral
diameter. The others are subuniformly about 0.23 or some-
what less. The turns commonly bulge in their middles.

TAXONOMY AND DISTRIBUTION

41

those of the mid-section more than the others. The wall is
rather thick, 0.06 oral diameter in places. There are thin
laminae with one layer of distinct radial rectangular prisms.
The wall is rather pallid.

Length, 15311.

The Carnegie lorica differs from that of Kofoid and
Campbell in that the spiral turns are right- rather than left-
turning; this is not due to error in drawing the reverse side.
Perhaps, as in some snails, right-wound forms of normally
left-turning shells may occur in Tintinnoina.

Coxliella pseudannulata differs from all the common
species of that genus in the shape of the aboral end. From
C. cdyptra it differs in the lack of pores or lacunae. How-
ever, calyptra is more likely one of the Nassellaria and not a
tintinnid. Coxliella pelagica lacks the contracted aboral horn
and median wide bulging turns, otherwise the two are
much alike. Coxliella decliuis has a narrower horn and
fewer spiral turns. Coxliella laciniosa, too, has a narrow
horn, and posterior lacunae.

Recorded from one station (10) in the Atlantic drift, in a
net sample taken at 50 meters. Frequency, 1 per cent.

Temperature, a?86; salinity, 34.94; density, 27.96; pH,
8.04.

CLIMACOCYLIS Jorgensen
Climacocylis, Kofoid and Campbell, 1929, p. 92.

Climacocylis, with its flaccid walls, open aboral end, and
distinctive form, is evidently specially adapted to life in the
warm seas where it is common. The genus is probably
derived from Coxliella.

Climacocylis is frequent in the warm waters near the
equator in both the Atlantic and the Pacific, and does not
venture far from this region to the north and south.

Three species are described here, and one new variety is
named.

Climacocylis scalaria (Brandt) Jorgensen

Climacocylis scalaria, Kofoid and Campbell, 1929, p. 93, rig.
155; Marshall, 1934, p. 640.

In its usual form the lorica consists of a long tube with a
spiral shelf and an expanded, flaring skirt; in length it is 6.9
to 10.0 oral diameters. The oral margin is entire and thin.
The long tubular section occupies about 0.59 the total
length, and on its upper two-thirds are 3 to 13 wide, plate-
like, left-turning spiral shelves. These shelves increase
gradually in diameter from oral to aboral end, ranging from
1. 1 8 to 1.92 oral diameters. The shelves occupy the middles
of wide spiral turns which number the same as the spiral
shelves, and which are unequal in width, the widest, at the
lower end, being 1.2 oral diameters. The shelves sometimes,
although not always, are connected by a jelly-like substance
recalling the outer sac of Brandtiella palliata. The lower-
most third of the tube is free of spiral turns and shelves, and
retains the same diameter as the oral end. At its aboral end
the squarish, widely flaring, flounced skirt arises. This skirt
has a length of about 2 oral diameters and is about as wide
as long. Its lower edges are incoherent and ragged. The

aboral end is wide open; in life it may be saccular and closed
off, but it is easily torn by the plankton net.

The wall is coarsely alveolar, with very large, subuniform
hexagons faintly visible in the soli, flaccid, and almost trans-
parent wall. There is but a single layer of prisms except in
the spiral shelves, which have three m seven layers. The
wall is subuniformly 0.01 oral diameter in thickness. Ad-
herent coccoliths, diatoms, and debris arc- not uncommonly
attached to the surface, sometimes extensively. The lumen
does not enter into the solid shelves.
Length. 24(1 to 344!<-

This peculiarly tropical species is exceedingly variable in
almost every character. One of the loricae from station 21
had a shape like a bottle with a wide, long neck. The whole
was spiraled and the shelves were not developed. The aboral
end was introverted.

Climacocylis scalaria suggests C. scalaroidcs, but is longer,
with hooplike skirt and wide, platelike shelves. A similar
spiral shelf occurs on Xystonella sea 11 dens, but that species is
altogether different; similar habits may have called forth
virtually identical structures in distant genera.

Recorded from forty-six stations, thirteen in the Atlantic
and thirty-three in the Pacific, as follows: two (2, 16) in the
Gulf Stream, five (17, 18, 19, 20, 21) in the Sargasso Sea,
four (22, 26, 27, 28) in the Atlantic equatorial region, two
(31, 34) in the Caribbean Sea. ten (40, 45, 46, 68, 69, 71, 73.
77, 78, 80) in the Galapagos region, one (54) in the South
Pacific middle latitudes, nine (82, 83, 84, 85, 87, 95, 157,
158, 159) in the region of South Pacific island fields, two
(99, 152) in the Pacific equatorial region, six (100, 109, 110-
in, 140, 150, 151) in the North Pacific trade region, three
(112, 113, 145) in the North Pacific middle latitudes, and
two (147, 149) in the California region. Climacocylis scalaria
avoids the Atlantic drift, North Sea, American cold-water
region, East Asiatic marginal sea, and Alaskan secondary
region. All these are regions of cool water. Its preference
for the warmer waters is also reflected in the temperature
records.

There are 17 pump and 49 net samples, of which 17 were
taken at the surface, 26 at 50 meters, and 23 at 100 meters.
Maximum frequency, 64 per cent at station 73 at the surface;
other records above minimum (2 to 51 per cent) from sta-
tions 2, 16, 17, 18, 19, 21, 34, 45, 46, 54, 71, 73, 77, 80, 82,
84, 85, 99, 109, iio-iii, 112, 113, 145, 147, 149, 150, 151,
157, 159. Thus, at thirty stations (45 per cent) this species
was one of the commoner Tintinnoina. It reached its max-
ima in the upper 50 meters, but did reach 2 per cent at 100
meters at station 54. Averages, 7.0 and 6.3 loricae in Atlantic
and Pacific pump samples, and 3.2 and 8.7 per cent in At-
lantic and Pacific net samples, respectively.

Temperature: Atlantic, pumpsamples i8?40-26?i9(23?89),
net samples i7?5o-26?57 (22?63); Pacific, i4?93-27?46
(23:64) and i5?33-28:74 (23^71 ), respectively. Salinity:
Atlantic, pump samples 36.08-37.15 (36.45), net samples
36.24-37.15 (36.61); Pacific, 33.68-36.49 (35-3°) and 34- 02 -
36.92 (35.67), respectively. Density: Atlantic, pump samples
23.95-26.01 (24.77), net samples 23.84-26.07 (25.16); Pacific,
21.60-25.98 (23.93) and 22.31-25.83 (23.92), respectively.

42

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

pH: Atlantic, pump samples 8.1 1-8.29 (8.22), net samples
8.16-8.32 (8.23); Pacific, 7.80-8.42 (8.18) and 7.87-8.39
(8.20), respectively.

Climacocylis scalaroides Kofoid and Campbell

Climacocylis scalaroides Kofoid and Campbell, 1929, pp. 93-94.
fig. 187; Marshall, 1934, pp. 641-642, fig. 14.

The short, fairly stocky lorica is finger-shaped and 2.6 to
6.0 oral diameters in length. The oral margin is smooth
and entire. The bowl has a collar-like spiral band of 3 to 17
left-wound turns of unequal width. These bands form
rounded ridges, and the collar-like section occupies 0.3 to 0.5
total length. The aboral five- to seven-tenths is devoid of
bands, and the whole tubelike lorica approximates a cone
(8°). The lateral contour is locally varied, especially in the
undivided region. The aboral end is commonly closed off
with some local thickening; sometimes it is open and ragged.

The soft, flaccid wall, easily deformed, has large hexagonal
prisms, about 65 around the mouth and 10 vertically across
the widest of the bands. The prisms are in a single layer
everywhere save in the suboral bands, where there may be
seven layers in the thickest parts. The wall is rather thick
in places, reaching nearly 0.3 oral diameter near the blobby
posterior part; mostly it is only a third as much.

Length, 90 to 271ft.

There is considerable variation in dimensions and propor-
tions, perhaps to be correlated with physical data. The num-
ber of spiral turns, their width, and the extent to which a
real shelf is developed are also characters subject to differ-
ence. Marshall (1934) found caudal flaps or tongues at the
aboral end; in the Carnegie material these tongues are lack-
ing, the loricae conforming to the originals. Specimens with
flaps may be distinctive in certain regions and should be
specially designated. The name suggested for such variety
is marshallae , n. var.

Close to Climacocylis scalaria in most characters, this
species differs, however, in being smaller, in having fewer
spiral turns in the collar, and in the lack of an aboral skirt,
the spreading flounces of which give a peculiar character to
scalaria. The spiral turns are never flat shelves as in scalaria,
and the aboral masses or flaps are different in the two. The
other species can hardly be confused with either of these,
although the tubular C. sip/10 has the spiral turns in a re-
stricted region; there is little else in common.

Recorded from seven stations, four in the Atlantic and
three in the Pacific, as follows: two (19, 21) in the Sargasso
Sea, one (22) in the Atlantic equatorial region, one (32) in
the Caribbean Sea, one (38) in the Pacific equatorial region,
and two (40, 73) in the Galapagos region.

There are 2 pump and 5 net samples, of which 3 were
taken at the surface, 3 at 50 meters, and 1 at 100 meters.
Frequency, 10 per cent at station 38; 2 per cent at station 40;
other records minimum.

Temperature: Atlantic, pump sample 19T21, net samples
i7?5o-26?57 (23? 12); Pacific, 28?oi and i5?33-26?48
(ao?9o), respectively. Salinity: Atlantic, pump sample 35.41,
net samples 36.28-37.15 (36.72); Pacific, 35.97 and 32.88-

34.89 (33.88), respectively. Density: Atlantic, pump sample
25.30, net samples 23.84-24.89 (24.36); Pacific, 23.14 and
21.31-25.83 (23.57), respectively. pH: Atlantic, pump sample
8.05, net samples 8.27-8.32 (8.29); Pacific, 8.23 and 7.87-8.33
(8.10), respectively.

Climacocylis sipho (Brandt) Kofoid and Campbell
Climacocylis sipho, Kofoid and Campbell, 1929, p. 94, fig. 184.

The lorica is an elongated tube with numerous spiral
turns, an open, ragged aboral end, and a length of 6.0 oral
diameters. The oral margin is thin and entire. The spiral
collar occupies the anterior 0.43 to 0.67 total length and is
made up of 29 to 33 left-wound, low-angled (5 to io°)
turns. The turns are subequal, although with a general tend-
ency to be wider in the lower part than in the upper. They
are separated from one another by optically dark, wide lines.
The lower 0.33 to 0.57 of the lorica, forming the bowl, is
tubular. The aboral region is devoid of spiral lines or other
special characters, the whole being really subconical (5 ).
The lateral contour is fairly regular. The wide-open aboral
end is either squarely truncated or ragged. In the latter case
it may be higher (as much as half the length of the aboral
section) on one side.

The wall is rigid and easily fractured. It is minutely
alveolar, with occasional lacunae in the spiral lamina.

There are 2 oval macronuclei and 18 membranelles.

Length, 260 to 290^1.

Climacocylis sipho is not close to other members of the
genus. Possibly it is closer to C. digitula or C. elongata than
to others. These species are similar in shape, but the spiral
is continuous from end to end. In this respect sipho is like
C. scalaria, but it has no expanded skirt and the wall is not
flaccid.

Recorded only from station 24 in the Atlantic equatorial
region, in a net sample taken at 50 meters. Frequency,
minimum.

Temperature, 23?i2; salinity, 36.00; density, 24.67; pH,
8.14.

Metacvlinae Kofoid and Campbell

Metacylineae Kofoid and Campbell, 1929, p. 197.

The Metacylinae include those genera in which the spiral
lamina is invariably limited to the anterior part of the lorica.
The subfamily was formerly assigned to the Petalotrichidae,
but is here transferred to the Coxliellidae, where it appears
to be more naturally placed.

The genera included are Metacylis and Hclicostomella.
Only the latter was found in the material of this expedition.

HELICOSTOMELLA ]6rgensen emended
Helicostomella, Kofoid and Campbell, 1929, p. 104.
Hclicostomella is related on the one hand to Coxliella in
having a spiral lamina, and on the other hand to Clima-
cocylis in having the spiral region limited. The wall has
typical secondary structure.

Hclicostomella is often common in temperate waters,
especially those of the north, although one species occurs as

TAXONOMY AND DISTRIBUTION

43

far south as Patagonia. Most of the species are well known
in northern Europe, and even to the north of Russia. Rarely
do species of this genus go far to sea, nearly all being within
coastal waters.

One species is described here.

Helicostomella longa ( Brandt) Jorgensen
Helicostomdla longa, Kofoid and Campbell, [929, p. 106, fig.
206.

The small, short, rather plump lorica, with slightly
swollen bowl and very short aboral spine, has a length of
2.24 oral diameters. The erect oral margin is formed by the
uppermost of the whorls that form the collar. The collar is
a spiral lamina of 4 whorls; its length is about 0.22 total
length, and the laminae are subequal in width. A fine, dark
line separates the turns one from another, and no overlap is
developed. The turns are laterally slightly concave. The
conical bowl is about 1.1 oral diameters in its greatest
diameter, which is reached near the middle. The lower part
narrows down (66°), and becomes more conical (90°)
nearer the tip. The aboral horn is about 0.2 oral diameter
in length, narrow conical (15°), and distally pointed.

The wall is finely alveolar, with tiny secondary Favella-
like hexagons. Collar and bowl are alike in this respect.
The wall is subuniformly 0.04 oral diameter in thickness,
with thin laminae, and has rectangular prisms in a single
layer. The lumen follows the contour exactly.

Length, 50 to 8op..

Helicostomella longa belies its name, being much the
shortest and plumpest member of the genus. Its collar has
fewer spiral turns than that of any other species, its bowl is
much the widest, and its horn is the shortest. It bears some
likeness to the antarctic Laac\manniella naviculijera, but
the bowl is pointed instead of open and lacks adherent
diatoms, and the collar has fewer turns.

Recorded from one station (130) in the California region,
in a net sample taken at 100 meters. Frequency, 42 per cent.

Temperature, %°.<)6\ salinity, 33.72; density, 26.14; pH,
8.06.

FAVELLIDAE Kofoid and Campbell emended

Favellineae, Kofoid and Campbell, 1929, p. 116.

In this family are included four genera, namely, Poroecus,
Favella, Protocymatocylis, and Cymatocylis. Only the first
two occur in the material of this expedition, the latter two
being antarctic. Poroecus is tropical, and Favella usually
coastal.

POROECUS Cleve emended

Poroecus, Kofoid and Campbell, 1929, pp. 116-117.

Poroecus is evidently the simplest member of the Favel-
lidae. Its thimble shape without much form differentiation,
entire to irregular oral margin, and wall structure are indi-
cations of this fact. The genus is possibly related to Tin-
tinnopsis.

Poroecus is uncommon in the tropical sea, where cocco-
liths are abundant.

One species is described here.

Poroecus apicatus Kofoid and Campbell
Poroecus apicatus Kofoid and Campbell, 1929, p. 118, fig. 223.

The small lorica, with subcylindrical upper and conical
lower bowl, has a length of 2.0 oral diameters. The oral
margin is irregularly ragged and not differentiated. The
bowl is cylindrical in the anterior half and is convex conical
(55°) in the posterior section. The aboral end is broadly
rounded.

The wall is thin and alveolar, and in each alveole there is
a coccolith that completely fills it.

Length, 56(1.

Poroecus apicatus is the shortest species of the genus. In
the lack of aboral horn it differs from all others save P.
brandti. From this species it differs in its shortness and con-
tracted lower bowl. In brandti the aboral end is hemispher-
ical and the lorica is more than twice as bulky, being 2.5
oral diameters and up to 2751.1 in length.

Recorded from one station (71) in the Galapagos region,
in a pump sample taken at 50 meters. Frequency, minimum.

Temperature, i8?85; salinity, 35.09; density, 25.86; pH,
7.83.

FAVELLA Jorgensen emended
Favella, Kofoid and Campbell, 1929, p. 147.

Favella agrees with Poroecus in having laminae, and in
similar wall structure and, in the simple species, similar
form. Favella is more advanced in having better finish,
larger size, and more highly developed form, and is also
related to Cymatocylis and Protocymatocylis. These last
two genera are exclusively antarctic in distribution, occurring
under the ice in the marginal region of the Antarctic Con-
tinent, off Kaiser William II Land, where they are probably
semineritic. No members of either of these genera are
recorded in the Humboldt Current, Cape Horn Current, or
Antarctic Current. The relation of these two genera to
Favella is clear both in form and in wall structure. Para-
javella is not related to any of these, but to the Xystonellidae.

Favella is almost exclusively found in neritic waters or in
water of coastal origin, where there are many local species.
A few species occur far out to sea, but these are generally
very different from coastal forms. As a whole, it is most
common in temperate water, but it does extend in range
from northern waters beyond Iceland to southward of Aus-
tralia. Some species cling to the coasts of tropical lands.

Five species are described here, of which two are new.

Favella adriatica (Imhof) Jorgensen emended Kofoid
and Campbell

Favella adriatica, Kofoid and Campbell, 1929, pp. 149-150, fig.

277.

The rather short, stout lorica, with narrow oral cuff, cup-
shaped bowl, and stout, conical aboral horn, has a length of
1.78 oral diameters. The oral margin is entire and a trifle

44

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

wavy. The erect oral cuff has a width of less than 0.04 oral
diameter and is quite hyaline. The cuplike bowl, with more
or less even upper margin, is narrow conical (4°) in the
anterior 0.78 oral diameter below the oral margin, and wider
(79 ) in the posterior 0.5 oral diameter. The sides are full
in the upper bowl and distinctly convex in the lower part.
The aboral horn is a cone (15 ) with a length of 0.5 oral
diameter. Its sides are sinuous and there are 2 or 3 short
pleats or folds near its upper end.

The wall is subuniformly thin, being not more than 0.02
oral diameter in thickness. The oral cuff is less than half as
much. There are well developed thin laminae and radial
prisms. The wall is everywhere finely meshed without
lacunae, and the mesh continues over the horn. The cavity
follows the outer contour.
Length, 200 to 400^1.

Favella adriatica has a shorter, stouter bowl than F. cam-
panula, as well as a wider horn. Its horn is shorter and less
spinelike than that of F. fislulicauda, and it has an oral cuff
lacking in that species. Its horn is better developed than
that of F. azorica, and that species has a relatively longer
bowl. Favella arcuata has a suboral ledge, a relatively longer
bowl, and a shorter horn. Favella injundibulum has a
conical instead of cuplike bowl and a handle-shaped horn.
Favella conjessa is of stouter facies, with a wider collar, a
serrated oral rim, and scarcely developed horn. Favella
attingata and F. serrata are very different species.

Recorded from one station (no) in the North Pacific
trade region, in a pump sample taken at 100 meters. Fre-
quency, minimum.

Temperature, 17^96; salinity, 34.73; density, 25.10; pH,
8.14.

Favella azorica (Cleve) Jorgensen
(Figure 56)
Favella azorica, Kofoid and Campbell, 1929, p. 151, fig. 284;

Marshall, 1934, p. 642, fig. 15.
The small, cuplike lorica, with no teeth, generally convex-
conical bowl, and aboral horn, has a length of 1.5 oral diam-
eters. The oral rim is entire, thin, and erect. There is no
differentiated collar, and the upper subcylindrical bowl con-
tinues with the same diameter as that of the oral opening for
a distance of 0.64 oral diameter. Its sides are full and evenly
contoured. The lower bowl contracts as a convex cone
(6o°) with a length of 0.57 oral diameter. The aboral horn
is a barely concave cone (55°) w ' tn a l en g tn of 0.39 oral
diameter, and it joins the lower bowl with gradual change
in contour.

The wall has a subuniform thickness of 0.1 oral diameter
across the bowl. It thins to the oral rim, and in the lower
bowl and horn is a little less. There are clear inner and
outer laminae, enclosing a single layer of radial, very faint
prisms. The wall shows no trace of annuli. It is glass-clear.
The animal has 2 oval macronuclei.
Length, 82U.

Marshall (1934) figures a lorica with suboral rings, alveolar
structure, and a length of 2 oral diameters. The length is
stated to be 73 to 107^1 and the oral diameter 47 to 65U. The

Carnegie loricae, which were abundant, show little signifi-
cant variation, all certainly lacking annuli.

Favella azorica most closely resembles F. composita, but
the latter species has a number of fine suboral rings and is
of somewhat different proportions. There is less difference
between bowl and horn in composita than in azorica. Favella
minutissima is tiny, with more sharply differentiated horn,
but does bear some general relationship to azorica. The
other species of Favella are unlike these species and can
hardly be confused with them.

Recorded from off Easter Island, in a net sample taken at
the surface. Frequency, 75 per cent; there were hundreds
of loricae.

No physical data were taken at this station.

Favella minutissima, new species
(Plate 1, figure 8; figure 59)
The exceedingly small, cup-shaped lorica has a length of
1.5 oral diameters. The oral margin is erect and simple.
There is no differentiated collar. The bowl continues below
the rim for about 1 oral diameter with the same diameter
throughout. Below this level it forms an inverted, rounded
subhemisphere. The aboral horn has a length of approxi-
mately 0.5 oral diameter, and is a simple, solid, pointed spike.
The wall is relatively thick, about 0.1 oral diameter, sub-
orally; toward the aboral end it becomes a little thinner. It
is hyaline, without traces of laminae, prismatic structure, or
surface markings of any sort. The cavity follows the outer
contour strictly, save that the horn is solid.

Length, total 28 to 3411, pedicel 3.1 to 6.211; oral diameter,
18.8 to 19U.

Favella minutissima differs from all others of the genus
in size, general proportions, and form. Its lack of collar and
of prismatic structure sets it off from others. If it had cocco-
liths it would have been placed with Poroecus, perhaps near
Poroecus apiculatus, but its size and shape are different. The
presence of a collar would have suggested Craterella, but it
lacks this essentially diagnostic organ. Its place in Favella is
uncertain, and it awaits final disposition. Favella azorica.
the species most like it in the genus, has no clear-cut horn.
Recorded from two stations (44, 75) in the Galapagos
region, in 1 pump and 1 net sample taken at 50 and 100
meters, respectively. Frequency, minimum.

Temperature: pump sample 20?g2, net sample i8?40.
Salinity: pump sample 34.86, net sample 35.47. Density:
pump sample 24.53, net sample 25.55. pH: pump sample
8.04, net sample 8.10.

Type locality, station 75, 100 meters; latitude 14 15'
south, longitude 92 05' west.

Favella septentrionalis, new species

(Plate 1, figure 12)

The moderately large, trim lorica has a narrow collar, a
long bowl, and a blunt, clumsy aboral horn; its length is
3.5 oral diameters. The narrow collar with median con-
striction is approximately 0.08 oral diameter in width. It is
outwardly concave, and its margin is slightly recurved

TAXONOMY AND DISTRIBUTION

45

toward the outside and inwardly convex. The collar is a
thin, membrane-like, hyaline cuff or ring with no trace of
spiral origin. Its upper margin is smooth, without irregu-
larity. The collar rests squarely upon the regular, neat bowl
below it. The bowl is long and thimble-shaped. The upper
part occupies 0.62 the total length. Its anterior diameter is
barely that of the oral opening, and it swells aborally to about
1. 1 7 oral diameters. This section, thus, forms a segment of
an inverted truncated cone (less than io°). Its sides are
even and smooth, and have no trace of even the slightest
irregularity. The aboral section of the bowl is somewhat
greater than a hemisphere, or, perhaps more correctly, re-
sembles half an oval. It occupies about 0.16 the total length;
its length is 1.75 of its own diameter at the oral end. Its
sides are full, even, regular arcs and like the upper section,
with which it is continuous. The blunt, thick aboral horn
has a length of about 0.22 total length. Its diameter at its
flaring base, where it joins the bowl, is 0.36 oral diameter,
and its free end is 0.42 basal diameter. It is, thus, also an
inverted cone (12 ). Its aboral end is almost squarely trun-
cated; its sides are sinuous and irregular, unlike the trim
lines of the upper parts.

The cavity of the lorica follows the outer contour closely
in the collar and bowl, but in the horn it is reduced to a very
narrow, slightly twisted, conical lumen which extends for
about 0.7 the length of the horn itself. The cavity is widest
orally and reaches a sharp point aborally. The wall is sub-
uniformly about 0.07 oral diameter in thickness across the
bowl, and 0.17 between the outer lamina and the wall of the
lumen of the pedicel. There are an inner and an outer
hyaline lamina, continuous about the oral end of the bowl.
Between these laminae occur the secondary prisms in a single
layer in the bowl. The outer lamina vanishes in the lower
section of the bowl, but the inner one continues to the lumen
of the pedicel, where it thins out and finally disappears. The
secondary prisms are larger, about 75 in number from oral
to lower end of the bowl. In the pedicel and also just be-
neath the collar the prisms occur in several layers, two or
three across below the collar, and four or five aborally. They
are less regular in shape in these regions. There are 3 to 45
prisms across the wall of the bowl. The aboral horn is with-
out striations or other surface marks, like the bowl.

Length, total 269(1, pedicel 57U; oral diameter, 7511.

Favella septentrionalis resembles F. jranciscana in general
facies, but has a thicker, blunter, clumsy horn instead of a
pointed one. The collar of jranciscana is a spiral, whereas
that of septentrionalis is a low ring. The wall of jranciscana
is roughened, unlike the trim one of septentrionalis. The
present species lacks the suboral bulge of F. arcuata and has
a different horn. It differs from F. serrata in having a
smooth, recurved collar and, again, a different horn. In
similar ways it differs from F. attingata. Favella ehrenbergii
lacks the collar of septentrionalis, and its horn is different.
Favella septentrionalis is more slender than F . conjessa and
of less square facies. It bears some resemblance to F . pana-
mensis, but that species has a relatively longer and more
pointed, regular horn and a taller collar, often with spiral
structure. The horn of septentrionalis is not at all like that

of F. campanula, which is also a trim lorica. It is propor-
tioned somewhat like F. tara\aensis, but the horn, again,
differs widely in form. The collar of tarakaensis is a spiral
and the margin is serrate.

Recorded from one station (116) in the North Pacific
middle latitudes, in 2 net samples, 1 each taken at the surface
and at 50 meters. Frequency, 4 per cent in both samples.

Temperature, ii?i8-i6?07 (13.62); salinity, 33.79-34.02
(33.90); density, 24.99-25.83 (25.41); pll, 8.11-8.17 (8.14).

Type locality, station 116, surface; latitude 38 41' north,
longitude 147 41' east.

Favella serrata (Mobius) Jorgensen
Favella serrata, Kofoid and Campbell, 1929, p. 156, fig. 293.

The generally cylindrical-conical, rather stout lorica, with
serrate oral rim, slight suboral bulge, and conical aboral
horn, has a length of 3.51 oral diameters. The oral margin
has about 70 minute, rounded triangular, subregular, slightly
incurved teeth. There is a low, hyaline, suberect cuff with
a width of less than 0.07 oral diameter immediately below
the margin, and the bowl contracts below it to a diameter
of 0.95 oral diameter at about 0.2 oral diameter below the
rim. Below this level it again swells out evenly to a diam-
eter of nearly 1.1 oral diameters at about 0.4 oral diameter
below the rim. This suboral bulge has a width of approxi-
mately 0.32 oral diameter. At the lower end the bowl
gradually and subregularly contracts as a cone (7 ) for a
length of nearly 1.6 oral diameters, and then as a wider one
(42 ) for 1.0 oral diameter. The upper section has more or
less full sides, and the lower cone is slightly convex. The
aboral horn is a narrow cone (io°) with a length of 0.86
oral diameter. Its sides are slightly sinuous and its tip is
pointed.

The wall has a subuniform thickness of not over 0.04 oral
diameter, and there are thin inner and outer laminae between
which there are thin-walled, radial, single layers of prisms.
The prisms are a little less wide suborally than in the lower
bowl. The surface displays a fine alveolar structure with an
occasional lacuna near the suboral part of the bowl. The
cavity of the lorica follows the outer contour except that the
horn is solid; the lumen enters into its upper end briefly.

The animal has 2 oval macronuclei.

Length, 320(1.

The Carnegie loricae are much more trim and less bulky,
with a relatively longer horn and with more pronounced
suboral contraction, than is typical. The horn is solid rather
than hollow.

Favella serrata differs from F. septentrionalis in having a
serrate oral rim and a thinner, less blunt aboral horn. Its
bowl is less conical than that of F. attingata and its horn is
longer. It lacks the alae of F. hclgolandica, and its bowl
and collar are different. Favella brcvis is squarish, with
suboral rings and a thicker horn. Favella conjessa has a
gradual transition between horn and bowl, and a very short
horn. Favella ehrenbergii lacks oral denticles, has no cuff
and no bulge, and has a wide, conical horn.

Recorded from two stations, one each in the Atlantic and

4 6

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

the Pacific, as follows: one (9) in the Atlantic drift, and one
(116) in the North Pacific middle latitudes.

There are 4 net samples, of which 2 were taken at the
surface and 2 at 50 meters. Maximum frequency, 8 per cent
at station 116; frequency at station 9, 2 per cent; averages,
1.5 and 5.5 per cent in the Atlantic and Pacific, respectively.

Temperature: Atlantic, 8?o6-i 1 : 12 ( 909); Pacific, 1 1 ? 1 8—
i6?07 (i3?62). Salinity: Atlantic, 35.11-35.14 (35.12);
Pacific, 33.79-34.02 (33.90). Density: Atlantic, 26.88-27.37
(27.12); Pacific, 24.99-25.83 (25.41). pH: Atlantic, 7.96-
8.u8 (8.02); Pacific, 8.11-8.17 (8.14).

PTYCHOCYLIDAE Kofoid and Campbell emended
Ptychocylidae, Kofoid and Campbell (part), 1929, p. 172.
Only the single genus Ptychocylis occurs in the family.

PTYCHOCYLIS Brandt emended Jorgensen
Ptychocylis, Kofoid and Campbell, 1929, pp. 185-186.

Ptychocylis is related in form and in wall structure to
Favella, Protocymatocylis, and Cymatocylis, and more re-
motely to Poroecus. Ptychocylis is the northern representa-
tive of these mostly southern and tropical genera.

Ptychocylis is distinctly a northern genus. Rarely does it
extend as far south as latitude 40 north. Mostly it occurs
in the Barents Sea, Davis Strait, Sea of Okhotsk, Bering
Sea, and north of the West Wind Current, often in minus
temperatures. Only washed and battered empty loricae enter
San Francisco Bay.

Six species are described here.

Ptychocylis acuta Brandt emended Kofoid and Campbell
Ptychocylis acuta, Kofoid and Campbell, 1929, p. 187, fig. 353.

The generally cylindrical lorica, with acutely conical aboral
end, has a length of 1.72 oral diameters. The oral margin is
fairly regularly serrate, with about 70 triangular, erect teeth
set upon a narrow, hyaline cuff with a width of only 0.03
oral diameter. The bowl swells concavely below the cuff to
a diameter of 1.17 oral diameters at 0.13 oral diameter below
the rim. This projecting ledge then rapidly narrows again
and there is a cylinder below it. A second ledge, with a
diameter of 1.17 oral diameters, is located at 0.45 oral diam-
eter below the rim. The lower part of the bowl is a segment
of an inverted truncated cone (25°) with a length of 0.77
oral diameter at its lower end, which is 1.22 oral diameters
below the rim; the diameter at this level is 0.83 oral diam-
eter. The aboral region forms a hollow-sided segment of an
inverted truncated cone (8o°) with a length of 0.25 oral
diameter. Its lower end is 1.47 oral diameters below the rim.
The diameter at this level reaches 0.38 oral diameter. The
distally blunted aboral horn is an inverted cone (50 ) with
a length of nearly 0.27 oral diameter.

The wall has a uniformly thin inner and outer lamina,
and clear interlaminar substance fills the space between. The
outer surface is irregularly reticulated with a scarcely raised
network of secondary structure.

Length, 120 to 145(1.

Ptychocylis acuta resembles P. wailesi in having an acute
aboral end, but the lower bowl is fuller and relatively shorter.
Ptychocylis glacialis has a sharply pointed aboral end, and
the remaining species are so different that they can hardly
be confused with acuta.

Recorded from one station (119) in the East Asiatic mar-
ginal sea, in a pump sample taken at the surface. There
were 2 loricae.

Temperature, 6:91; salinity, 32.96; density, 25.85; pH,
7.96.

Ptychocylis arctica Brandt emended Kofoid and Campbell
Ptychocylis arctica, Kofoid and Campbell, 1920, p. 187, fig. 351.

The large, stout, bullet-shaped lorica, with blunted, flat-
tened aboral end, has a length of 1.48 oral diameters. The
oral margin has about 50 large, triangular, erect teeth, evenly
spaced and subequal. These denticles arise from a hyaline
cuff with a width of only 0.03 oral diameter. Below the cuff
the bowl extends as a narrow cylinder for a width of 0.05
oral diameter, and an angular ledge with a width of 0.06
oral diameter and a diameter of 1.12 oral diameters occurs
below this cylinder. The bowl extends as a cylinder below
the ledge once more, and is 0.25 oral diameter in width. At
its lower end is a second ledge with a width somewhat less
than that of the upper one, but with the same diameter. The
lower part of the bowl contracts as an inverted truncated
segment of a cone (22°) with a width of 0.78 oral diameter;
its sides are plane to slightly full. The aboral region is an
inverted convex cone (83 ) with a width of 0.26 oral diam-
eter. The aboral end is squarely truncated, and its diameter
is 0.19 oral diameter.

The wall is thickened in the ledges to 0.05 oral diameter,
and is less than half as thick in the lateral region. Laminae
are difficult to detect, and the wall is homogeneous. The
outer surface is reticulated with rather small prisms, the
edges of which are weakly raised, giving the surface, espe-
cially at the aboral end, a pitted structure. The clear cuff
rests upon the dense bowl. The lumen does not enter the
ledges.

Length, 120 to 140(1.

Ptychocylis arctica differs from P. drygalsk^u in the longer,
lower bowl and general bullet shape. From P. basicurvata
it differs in aboral truncation rather than simple aboral
rounding. The remaining species have altogether different
aboral ends.

Recorded from one station (10) in the Atlantic drift, in 2
net samples, 1 taken at the surface and 1 at 50 meters. Fre-
quency, minimum.

Temperature, 9?86-io?94 (io?4o); salinity, 34. 94-34- <is
(34.945); density, 26.77-27.96 (27.36); pH, 8.04-8.08 (8.06).

Ptychocylis drygalskii Brandt

Ptychocylis drygalskii, Kofoid and Campbell, 1929, p. 188, fig.
350; Hada, 1932^, p. 567, fig. 11.

The heavy-set lorica, with little-developed ledges and
truncate conical, flattened aboral end, has a length of 1.4
oral diameters. The oral margin is regularly provided with

TAXONOMY AND DISTRIBUTION

47

about 34 erect, triangular teeth. These teeth arise from a
narrow (0.02 oral diameter) hyaline cuff. The bowl below
the cuff expands concavely, giving rise to an angular ledge
of nearly 1.15 oral diameters, close to 0.15 oral diameter
below the rim. The bowl below the ledge is concave
cylindrical. This cylindrical part is nearly 0.4s oral diameter
in length, and at its lower end there is a ledgelike swollen
/one of about 1.2 oral diameters with a width of approxi-
mately 0.22 oral diameter; sometimes this zone is more
sharply marked and not so wide. The bowl contracts below
this second ledge as an inverted basal segment of a hollow-
sided truncate cone (27 ) with a diameter, at its lower end,
of 0.94 oral diameter. The aboral region again contracts as
a concave cone (76 ) with a width of 0.=; oral diameter, and
the broadly rounded aboral end has a diameter of nearly
0.25 oral diameter.

The wall is homogeneous, and its structure is difficult to
make out. Its thickness hardly exceeds 0.01 oral diameter,
even in the ledges. The cavity partially enters the ledges
and the horn. The surface shows hexagonal meshwork and
is weakly pitted. It is denser aborally than elsewhere.

Length, 77U..

The Carnegie loricae are more like those of Hada (19326)
than those of Brandt in size, proportions, and other charac-
ters; perhaps there is some geographical difference.

Ptychocylis drygalskji resembles P. obtusa, but it is stouter
and its aboral end is not similar. It bears some likeness to
P. arctica, but is shorter and stouter and has a less bullet-
shaped bowl than that species; arctica has a narrower conical
aboral region and pointed tip.

Recorded from one station (123) in the East Asiatic mar-
ginal sea, in a pump sample taken at the surface. Frequency,
minimum.

Temperature, 8? 10; salinity, 32.76; density, 25.52; pH,
8.03.

Ptychocylis minor Jorgensen
Ptychocylis minor, Kofoid and Campbell, 1929, p. 188, fig. 354.

The rather short, urn-shaped lorica, with conical aboral
region and short horn, has a length of 1.7 oral diameters.
The oral margin is regularly provided with about 56 short,
erect, bluntly pointed, evenly spaced, triangular teeth resting
on a narrow, hyaline cuff, the width of which is only 0.01
oral diameter. The bowl expands (8o°) to a narrow, pro-
jecting shelf or ledge, the diameter of which is 1.18 oral
diameters, and which is at 0.11 oral diameter below the rim.
Below this shelf the bowl contracts with concave lateral con-
tour to a swollen zone (1.08 oral diameter) with a width of
0.22 oral diameter, near 0.5 oral diameter below the rim.
Below this swelling the bowl contracts as an inverted trun-
cated, barely concave cone (16°) with a width of 0.39 oral
diameter, and a diameter at its lower end of 0.92 oral diam-
eter. The aboral region is an inverted hollow cone (90 ),
with a width equal to that of the region just above it, and a
diameter of 0.21 oral diameter at the lower end. The aboral
horn is a narrow cone (30°), with a length of 0.34 oral
diameter. Its tip is minutely rounded.

The wall has thin laminae, and two to five layers of tiny,

hexagonal prisms are thus enclosed. The wall is thickest
suborally and in the horn, where it reaches nearly 0.05 oral
diameter; elsewhere it is about hall as thick. The outer sur-
face is rather coarsely reticulated with pitted hexagons; these
pits are especially developed in the shell and aboral cone.

Length, 90 to 13511.

Some of the Carnegie loricae contract with sharp angles
suborally and have decided conical bowls and bluntly conical
horns.

Ptychocylis minor most resembles /'. urnula, but is shorter,
with less conical aboral region and more conical bowl. The
horn is also shorter and more conical. Ptychocylis acuta has
a shorter horn; P. repanda and P. ostenjeldi are similar.

Recorded from seven stations in the Atlantic, as follows:
one (7) in the North Sea, four (8, 9, 10, 11) in the Atlantic-
drift, one (12) in the American cold-water region, and one
(14) at the margin of the Gulf Stream.

There are 2 pump and 14 net samples, of which 6 were
taken at the surface, 6 at 50 meters, and 4 at 100 meters.
Maximum frequency, 100 per cent at station 10 at the sur-
face; other records above minimum (2 to 80 per cent) from
stations 7, 8, 9, n, 12, 14; averages, 5.5 loricae and 53.2 per
cent in pump and net samples, respectively.

Temperature: pump samples 7?oi-8?44 (7^72), net
samples 4?i8-i4?02 (8^96). Salinity: pump samples 33-65-
34.97 (34.31), net samples 34.51-35-59 v35-°2)- Density:
pump samples 26.16-27.42 (26.79), net samples 26.66-27.96
(26.96). pH: pump samples 7.92-8.10 (8.01), net samples
7.92-8.08 (8.00).

Ptychocylis obtusa Brandt emended Kofoid and Campbell

(Figures 57, 58)

Ptychocylis obtusa, Kofoid and Campbell, 1929, pp. 188-189,
fig. 349; Schulz and Wulff, 1929, pp. 336-338, figs. 105-109;
Bernstein (part), 1931, pp. 13-14, fig. 2a, b (for fig. zc-g see
P. glacialis and P. urnula); Hada, 1932^, pp. 55-56, fig. 21;
19326, p. 567.

The stout, squat lorica, with aboral cone and flattened end,
has a length of 1.57 oral diameters. The oral margin is
regularly serrate with minute, evenly spaced triangular
points, and these rest on a very narrow, clear band the width
of which is only 0.02 oral diameter. The bowl expands con-
cavely and conically (60°) for a distance of 0.14 oral diam-
eter and reaches 1.13 oral diameters at its lower end. It
again contracts conically (50 ) and reaches a diameter of
1.04 oral diameters at 0.26 oral diameter below the rim.
Once again it expands conically (60°), reaching a diameter
of 1.22 oral diameters near 0.43 oral diameter below the rim.
Below this level the bowl contracts conically (65°), reaching,
at 0.55 oral diameter below the rim, a diameter of 1.07 oral
diameters. This last diameter is maintained for a distance
of 1. 12 oral diameters below the rim, giving the central
region of the bowl a generally cylindrical contour. At the
lower end of this cylinder the bowl contracts conically (8o°)
once more for 0.24 oral diameter, reaching a diameter of
0.68 oral diameter at the lower end of this zone. The pedicel-
like region is a truncated cone (45°) with a length of 0.43

4 8

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

oral diameter. Its lower end is squarely truncated and its
width is 0.4 oral diameter. The sides of this much contracted
lorica are neatly rounded over the several shoulders, and the
conical regions are generally full or convex.

The wall is usually thin, hardly exceeding 0.01 oral diam-
eter, with some slight thickenings at the angles. Laminae
and alveolar structure are not distinguished. The outer
surface is reticulated, however, with elongate hexagons, and
near the aboral end it is pitted, the walls of the reticulations
being lifted above the general surface. The lorica is dark
and heavy.

Length, 119(1.

These extremely numerous loricae are decidedly variable.
One extreme form is shown (fig. 58). Schulz and Wulff
(1929) illustrate a long series taken in different temperatures
and other physical conditions in the Barents Sea; these loricae
all differ widely.

Ptychocylis obtusa resembles P. cylindrica, but the cylin-
drical part of the bowl is not so long and the lorica is much
stockier. The character of the aboral end differentiates obtusa
from the other species, for none of them has a wide, flat
end; their ends are, for the most part, either acute or pointed.

Bernstein (1931) included P. glacialis and P. urnula in
obtusa. Some of Schulz and Wulff's loricae (1929) belong
elsewhere (especially in P. cylindrica), although not distin-
guished here. The genus needs special study.

Recorded from thirteen stations, three in the Atlantic and
ten in tht Pacific, as follows: one (7) in the North Sea, one
(10) in the Atlantic drift, one (13) in the American cold-
water region, two (116, 117) in the North Pacific middle
latitudes, six (118, 119, 120, 121, 122, 123) in the East Asiatic
marginal sea, and two (124, 125) in the Alaskan secondary
region. The species is distinctly circumpolar.

There are 18 pump and 21 net samples, of which 10 were
taken at the surface, 15 at 50 meters, and 14 at 100 meters.
Maximum frequency, 91 per cent at station 125; other records
above minimum (2 to 79 per cent) from stations 13, 116,
117, 120, 121, 122, 123, 124. Records of 2 to 300 loricae
occur in pump samples; average frequencies in net samples,
15 and 44.2 per cent in Atlantic and Pacific, respectively.

Temperature: Atlantic, pump sample n?27, net samples
1? 64-11? 27 (7^72); Pacific, i?72-io?50 (5?og) and 2?oi-
i8?i8 (5?8o), respectively. Salinity: Atlantic, pump sample
32.68, net samples 32.68-35.24 (34.06); Pacific, 32.73-33.72
(33.00) and 32.73-34.06 (33.12), respectively. Density: At-
lantic, pump sample 24.94, net samples 24.94-27.96 (26.81);
Pacific, 25.13-26.52 (25.48) and 25.52-26.76 (26.16), respec-
tively. pH: Atlantic, net samples 8.04; Pacific, pump samples
7.86-8.21 (7.95), net samples 7.64-8.1 1 (7.97). Records of
pH were not available for the Atlantic pump sample, but
were available for 2 of the net samples, each 8.04.

Ptychocylis urnula (Claparede and Lachmann) Brandt
Ptychocylis urnula. Kofoid and Campbell, 1929, pp. 189-190,
fig. 355; Schulz and Wulff, 1929, pp. 336-338, figs. 111-114.
Ptychocylis obtusa, Bernstein (part), 1931, pp. 13-14. rig. 2/, g

(for fig. la-e see P. obtusa and P. glacialis) .
The generally cylindrical lorica, with conical aboral end

and elongated, pointed horn, has a length of 1.93 oral diam-
eters. The oral margin is regularly set with about 60 sharp,
erect, evenly spaced, triangular teeth, which arise from a
hyaline cuff with a width of less than 0.02 oral diameter.
The bowl swells concavely to a suboral ledge of 1.17 oral
diameters at 0.1 oral diameter below the rim. Below the
ledge it assumes a barely concave cylindrical shape for a
distance of 0.37 oral diameter below the rim, at which level
a second, less pronounced and distinctly rounded, ledge
occurs. This ledge has a diameter of 1.06 oral diameters and
occupies a zone with a width of approximately 0.14 oral
diameter. The bowl below this zone becomes subcylindrical
for 1.0 oral diameter and again swells slightly, reaching a
diameter a little greater than that of the oral margin. Below
this level the bowl contracts as a plane or barely concave
cone (62 ) with a length of 0.68 oral diameter. At its lower
end is the aboral horn, which is a narrow, sharp, pointed
cone (26 ) with a length of nearly 0.26 oral diameter.

The wall is thick, being 0.06 oral diameter across the
upper ledge, where it is thickest; elsewhere it is only about
a third as much. The cavity follows the outer contour, enter-
ing into the upper bulge and extending to the tip of the
horn. The wall is denser aborally and its surface is finely
reticulated with irregularly formed hexagonal prisms, the
walls of which are raised so that the bowl appears pitted.

Length, 123 to 192^.

Variations in length are probably correlated with tem-
perature. Some loricae have longer horns than others, but
the general shape is retained.

Ptychocylis urnula resembles P. minor, but is longer and
has a more conical aboral region and a thinner and longer
horn. Its bowl is also more cylindrical. Its horn is not so
long as that of P. repanda or P. ostenfeldi, and the bowls of
these two species are very different. Ptychocylis wailesi has
a concave-conical lower bowl and lacks a prolonged horn.
Other species have rounded aboral ends and are not closely
related.

Bernstein ( 1 93 1 ) unites urnula and obtusa in his figures,
but distinguishes one lorica in his text as urnula (fig. 2g).

Recorded from one station (10) in the Atlantic drift, in a
net sample taken at 100 meters. Frequency, minimum.

Temperature, 6^56; salinity, 35.02; density, 27.52; pH,
7-95-

PETALOTRICHIDAE Kofoid and Campbell emended
Petalotrichidae, Kofoid and Campbell, 1929, p. 190.
Included in this family are three genera: Craterella, Acan-
thostomella, and Petalotricha. All three genera occur in the
material of this expedition. Petalotricha is exclusively a
warm-water genus, the others being more widely spread,
often into very cold waters.

Craterellinae Kofoid and Campbell
Craterellineae Kofoid and Campbell, 1929, p. 190.
Two genera belong to this subfamily: AcanthostotneUa

and Craterella.

TAXONOMY AND DISTRIBUTION

49

CRATERELLA Kofoid and Campbell
Craterella Kofoid and Campbell, 1929, p. 194-

Craterella includes a few small tropical species as well as
one or two from cooler waters. Owing to the small dimen-
sions, the actual distribution is inadequately known, as are
the species.

Two species are described here.

Craterella armilla Kofoid and Campbell
(Figure 19)
Craterella armilla Kofoid and Campbell, 1929, p. 195, fig. 371.

The tiny, saccular lorica, with low inner collar, repeated
outer collars, thimble-like bowl, and pointed aboral end, has
a length of 1.53 oral diameters. The oral margin is thin,
erect, and entire, and the inner collar spreads to the upper
end of the anteriormost of the two outer collars as a segment
of a cone (62 ) with a length of hardly 0.08 oral diameter.
The upper outer collar has a diameter of 1.15 oral diameters,
and flares (45°). It is separated by a trough from the inner
collar. The second outer collar has a diameter of 1.3 oral
diameters, and flares (90 ). It is separated from the first
outer collar by a trough. The pointed bowl has a length of
1.38 oral diameters. Its upper two-thirds is a truncated seg-
ment of an inverted cone (10°), and it has plane sides. The
aboral third of the bowl is conical (8o°), with convex sides
and a simple pointed aboral end.

The wall is thick, with distinct laminae and interlaminar
prisms, and the surface shows rugosity.

Length, 29U,.

The lorica figured (fig. 19) has a pointed rather than
rounded aboral end.

Craterella armilla differs from the other species in having
repeated collars and a rough surface. It is stouter than C.
protuberant and less contracted below. In general shape it
is like C. torulata, but the collar duplication differentiates it.
The collars flare instead of contracting as in C. urceolata.
Craterella aperta from Australia is not at all like armilla, or,
indeed, any of the others; it may not belong to Craterella.

Recorded from two stations, one in the Atlantic and one
in the Pacific, as follows: one (23) in the Atlantic equatorial
region, and one (69) in the Galapagos region.

There are 1 pump and 1 net sample, taken at the surface
and at 50 meters, respectively. Frequency, minimum. The
pump sample was taken in the Atlantic and the net sample
in the Pacific.

Temperature: pump sample 21 ? 1 3, net sample 20?gg.
Salinity: pump sample 35.24, net sample 36.04. Density:
pump sample 24.66, net sample 25.30. pH: pump sample
8.12, net sample 8.14.

Craterella urceolata (Ostenfeld) Kofoid and Campbell

(Figure 20)
Craterella urceolata, Kofoid and Campbell. 1929. p. 196, fig. 368.

The tiny lorica has a contracted aperture, flaring outer
collar, and wide, conical, pointed bowl, and its length is 1.24

oral diameters. The oral margin is thin, sharp-edged, and
entire. The inner collar is a basal segment of a truncated
cone (8o°) with concave sides and a trough separating it
from the outer collar below. The outer collar has a diameter
of 1.2 oral diameters and a width of 0.16 oral diameter. It
flares (54°) from the bowl below, the upper diameter of
which is 1.1 1 oral diameters. The upper two-thirds of the
bowl is a segment of an inverted truncated cone (30 ), ami
the lower section is a wider cone (82 ). The sides of both
sections are convex, those of the upper section less so than
those of the lower. The aboral end is pointed but not pro-
longed or sharp.

The wall is hyaline and thin, ami the cavity neatly follows
the outer contour.

Length, 48^1.

The lorica figured (fig. 20) has a more conical bowl, more
flare in the outer collar, and a narrower aperture than usual.

Craterella urceolata differs from other species in having a
high inner collar, and in its stouter proportions. It is some-
what like C. obscura, but the bowl is not so long, and its
upper section is less cylindrical. It bears some likeness to
C. acuta, but that species has a higher, more flaring inner
collar, and a pointed, prolonged aboral end. Craterella pro-
tuberant has a blunt aboral horn.

Recorded from one station (41) in the Galapagos region,
in a net sample taken at the surface. Frequency, minimum.

Temperature, 20?42; salinity, 34.19; density, 24.06; pH,
8.11.

ACANTHOSTOMELLA Jorgensen

Acanthostomella, Kofoid and Campbell, 1929, p. 191.

Acanthostomella adds a number of teeth to the outer collar
of Craterella. The genus is, however, remarkable in that
there are two sharp divisions in distribution. One group of
species is distinctly tropical, and the other group is of cold-
water facies. The latter mostly have distinct prismatic struc-
ture, whereas the former are glass-clear. The tropical species
usually have fewer and stronger teeth than the others. As
yet the genus is not so well known as it will be on further
examination of microplankton.

Six species are described here.

Acanthostomella elongata Kofoid and Campbell
Acanthostomella elongata Kofoid and Campbell, 1929, p. 192,
fig- 359-

The tall lorica is shaped like a finger cot and has a sub-
hemispherical aboral end and a short spine; the length is 3.0
oral diameters. The oral margin is thin and entire, and the
inner collar is erect and rather wide. The outer collar has
24 outward-directed, low clawlike teeth, and hardly flares.
The upper bowl is cylindrical and about 2.6 oral diameters
in length. The lower bowl is about 0.4 oral diameter in
length, or less, and is an inverted convex cone (62 ). Its
lower end is drawn out into a tiny aboral spinule.

The wall is thin: alveoles and laminae are present.

Length, 7011.

Acanthostomella elongata has a long cylindrical bowl, at

5°

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

least twice as long as that of any other species. It resembles
a small replica of Parafauella digitalis, but that species not
only is larger and longer, but also lacks teeth and inner and
outer collars. It is not at all like the common A. norvegica,
its frequent neighbor.

Recorded from one station (125) in the Alaskan secondary
region, in a pump sample taken at 50 meters. There were
2 loricae.

Temperature, 5?68; salinity, 32.79: density, 25.88; pH,
7.98.

Acanthostomella gracilis (Brandt) Kofoid and Campbell

Acanthostomella gracilis, Kofoid and Campbell, 1929, p. 192,
fig. 360.

The rather short, stout lorica, with cylindrical upper bowl
and conical, pointed aboral end, has a length of 1.7 oral
diameters. The oral margin is thin and erect; the inner
collar is cufflike and its width is nearly 0.07 oral diameter.
It is separated by an angular trough from the outer collar.
The outer collar flares (35°) and from its free edge arise 36
narrow, outward-projecting, subequal, subequidistant, tri-
angular teeth. The upper bowl is cylindrical and about 1.1 1
oral diameters in length. The bowl is conical (80°) with
distinctly convex sides, and its lower end is drawn out
bluntly.

The wall is rather thick, about 0.03 oral diameter in thick-
ness, and there are thin laminae with rectangular, radial
secondary prisms enclosed between them. The wall thins
down a bit at the oral end.

The large animal has 2 macronuclei and rather long mem-
branelles.

Length, 45 to 6o|x.

Acanthostomella gracilis has a cylindrical upper bowl and
a bluntly pointed aboral end, whereas A. norvegica, its closest
relative, has a laterally concave and aborally swollen bowl
with a distinct horn. Acanthostomella obtusa has a blunted,
distinct, and longer horn, and A. elongata has a much taller
bowl.

Recorded from one station (119) in the East Asiatic mar-
ginal sea, in 2 pump samples taken at 50 and 100 meters.
Frequency, 2 and 4 loricae.

Temperature, 3?42 at 50 meters; salinity, 33.05-33.13
(33.09); density, 26.31-26.52 (26.41); pH, 7.85-7.93 (7.89).

Acanthostomella lata Kofoid and Campbell
Acanthostomella lata Kofoid and Campbell, 1929, pp. 192-193.

fig- 364-
The stout, goblet-shaped lorica, with low teeth, high inner
collar, conical bowl, and sharp, pointed aboral spine, has a
length of 1.25 oral diameters. The oral margin is thin, erect,
and entire. The inner collar is cufflike and about 0.1 oral
diameter in width. The outer collar, which is separated
from the inner by a deep trough, flares (20 ) and its rim
has 22 short, stout, outward-spread triangular teeth whose
free tips are rotated to the left. The bowl is conical (about
15 ) suborally, and wider conical (90°) toward the lower
end. It is full and distinctly convex. The aboral horn is a

spine (20°) with a length of approximately 0.25 oral diameter.

The subuniformly thin wall has clearly marked laminae
and distinct prismatic structure. The spine is hollow.

Length, 3711.

The Carnegie loricae agree closely with that of Kofoid
and Campbell, save that the wall is unusually glassy.

Acanthostomella lata is larger than most species, and rela-
tively wider, with shorter teeth and higher inner collar. It
has more teeth than A. conicoides, a distinctly convex instead
of concave lower bowl, and a distinct spine. Its bowl is less
saccular than that of the northern A. norvegica, its spine is
longer, and its teeth are shorter, fewer, and rotated.

Recorded from five stations, one in the Atlantic and four
in the Pacific, as follows: one (33) in the Caribbean Sea, one
(40) in the Galapagos region, one (65) in the South Pacific
middle latitudes, one (137) in the California region, and one
( 159) in the region of South Pacific island fields.

There are 1 pump and 4 net samples, of which 1 (137)
was taken at the surface and 2 each at 50 and 100 meters.
Frequencies did not exceed 1 per cent.

Temperature: Atlantic, net sample 23?^; Pacific, net
samples i5?03-27?oo (i9?42), pump sample 25?48. Salinity:
Atlantic, net sample 36.49; Pacific, net samples 34.30-35.75
(34.98), pump sample 34.97. Density: Atlantic, net sample
25.03; Pacific, net samples 23.01-25.83 (24.75), pump sample
23.19. pH: Atlantic, net sample 8.18; Pacific, net samples
7.83-8.37 (8.10), pump sample 8.39.

Acanthostomella minutissima Kofoid and Campbell

Acanthostomella minutissima Kofoid and Campbell, 1929, p.
193. fig- 358.

The rather plump, short, cuplike lorica, with numerous
teeth and round aboral end, has a length of 1.30 to 1.64 oral
diameters. The oral margin is thin and erect. The inner
collar is narrow, cufflike, and thin. The outer collar flares
sharply (50 ) and carries about 25 slender, narrow triangu-
lar, outward-curved teeth which are about as long as the
width of the inner collar. The bowl is subcylindrical in the
anterior 0.55 and contracts as a cone (90°) in the lower
section. This aboral region is subhemispherical, with con-
vex sides. The aboral end is not especially differentiated.

The rather thick wall has thin laminae and is pronouncedly
alveolar. Kofoid and Campbell (1929) found coccoliths
studding the whole exterior of some loricae. The cavity con-
forms to the outer contour.

Length, 29 to 36^.

The shape of the aboral end is hemispheroidal to parabo-
loidal, and the wall may have coccoliths.

This small tropical species generally resembles the north-
ern Acanthostomella norvegica, but differs in being smaller,
and in having a distinct, flaring collar. Its upper bowl is
typically not so long and the aboral end never has the
emergent point of norvegica. It is unlike any of the other
tropical species. Had it no teeth it would have been con-
sidered something like Cratcrella urceolata.

Recorded from four stations, one in the Atlantic and three
in the Pacific, as follows: one (18) in the Sargasso Sea, two

TAXONOMY AND DISTRIBUTION

51

(41, 69) in the Galapagos region, and one (65) in the South
Pacific middle latitudes.

There are 4 net samples, of which 2 were taken at the
surface and 2 at 100 meters. Frequency, 2 per cent at
station 69.

Temperature: Atlantic, 20?32; Pacific, i5?oo-2i? 13 (i8?86).

Salinity: Atlantic, 36.81; Pacific, 34.19-3s.24 (34.57). Den-
sity: Atlantic, 26.01; Pacific, 24.06-25.44 (24.72). pH: At-
lantic. 8.21; Pacific, 8.10-8.12 (8.11).

Acanthostomella norvegica (Daday) Jorgensen

(Figures 21, 22, 23)

Acanthostomella norvegica, Kofoid and Campbell, 1929, p. 193,
fig. 363; Bernstein, 1931, p. 14; Hada, 1932,;. pp. 56-57. fig.
22; 19326, p. 567, fig. 20.

The short, stout, cuplike lorica, with short, incurved, claw-
like teeth, wide, swollen bowl, and aboral end with tiny
nipple, has a length of 1.14 to 1.71 oral diameters. The oral
margin is thin. The inner collar is less than 0.1 oral diameter
in width, and has a spreading base. There is a trough be-
tween it and the outer collar. The outer collar flares (40 to
50 ) and has outwardly convex sides. On its free edge are
20 to 36 incurved, clawlike, sharply pointed, narrow tri-
angular, subequal, and subequidistant teeth. The upper
section of the short, wide bowl is an inverted basal segment
of a full truncated cone (4 ) with a length of 0.73 to 1.0
oral diameter. The lower section is at first 42 to 63 , and
later 90 to 145 , thus forming a rounded cone, or subhemi-
sphere, with pronouncedly convex sides. The aboral end has
a nipple-like horn with a length of not more than 0.1 oral
diameter.

The wall is uniformly thin (hardly 0.02 oral diameter).
There are laminae with rectangular, subequal prisms in a
single layer. The inner collar is glass-clear. The horn is
solid. The aboral half of the bowl is mostly plastered with
adherent fecal matter.

There are 2 oval macronuclei.

Length, 45 to 50U..

Three extreme variants are shown (figs. 21-23). Of these,
one (fig. 23) is hollow-sided in the upper bowl, and swollen
at about 1 oral diameter below the rim. Its bowl is longer
than usual. The others figured are more nearly of the usual
type. Hada (1932a, 19326) figures 2 loricae, one with a
longer horn than the other. The horns on these specimens
are longer and more sharply pointed than those on the
Carnegie specimens. The bowls are not so long as that of
the extreme specimen of this collection, nor so short as those
of others. In other characters, however, all these agree.

Acanthostomella norvegica is, perhaps, closest to A.
gracilis, which has the same general habitat. It is aborally
expanded, however, and has a nipple. Its lower bowl is
conical rather than cylindrical, and the aboral region is less
angular. Its bowl is not so long as that of A. elongata, and
it is altogether unlike the tropical species of the genus in
most characters.

Recorded from fifteen stations, eight in the Atlantic and
seven in the Pacific, as follows: one (7) in the North Sea,

three (8, 10, 11) in the Atlantic drift, two (12, 1 \) in the
American cold-water region, one (14) in the Gulf Stream,
two (117, 126) in the North Pacific middle latitudes, four
(120, 121, 122, 123) in the East Asiatic marginal sea, one
(124) in the Alaskan secondary region, and one (130) in
the California region. Acanthostomella norvegica has a pro-
nounced northern and temperate distribution and appears
elsewhere either accidentally in watei ol warmer origin, as
in the Gulf Stream (station 14), or as a subsurface dweller.

There are 14 pump and i(> net samples, of which 9 were
taken at the surface, 13 at 50 meters, ami 8 at 100 meters.
Maximum frequency, 68 per cent at station 13; other records
above minimum (2 to 56 per cent) from stations 7, 11, 14,
117, 120, 121, 123, the greatest frequencies being in surface
catches; averages, 19.0 and 1.8 loricae in Atlantic and Pacific
pump samples, and 14.8 and 7.5 per cent in Atlantic and
Pacific net samples, respectively.

Temperature: Atlantic, net samples 4?i8-i4?02 (8^55),
pump samples 7?oi-i4?95 (n?07); Pacific, 2?i7-i2?9i
(9? 10) and 2?05~i2?56 (7? 18), respectively. One record
of i4?02 was in the Gulf Stream, where 3 empty loricae
were recorded in 2 samples. Salinity: Atlantic, net samples
32.68-35.95 (34.73), pump samples 32.68-35.10 (34.25);
Pacific, 33.06-34.22 (33.60) and 32.66-34.22 (33.07), respec-
tively. Density: Atlantic, net samples 24.94-27.96 (27.04),
pump samples 24.94-27.42 (26.14); Pacific, 25.19-26.41
(26.00) and 25.26-26.48 (25.84), respectively. pH: Atlantic,
net samples 7.93-8.08 (8.01), pump samples 7.92-8.18 (8.05);
Pacific, 7.72-8.26 (7.99) and 7.84-8.06 (7.98), respectively.

Acanthostomella obtusa Kofoid and Campbell

Acanthostomella obtusa Kofoid and Campbell, 1929, p. 194,
fig- 361.

The short, stout, cuplike lorica, with tall, thickened inner
collar, heavy outer collar, few teeth, and short, blunt aboral
horn, has a length of 1.6 to 1.7 oral diameters. The oral
margin is thin, erect, and entire, and the wider outer collar
has 20 short, stout, outward-projecting, triangular teeth.
The outer collar is less than 1.1 oral diameters in diameter
and barely flares. The upper bowl is a full truncated cone
(20 ), and the lower three-tenths widens out as a hollow
cone (68°). The lower part of the aboral three-tenths widens
yet more. The aboral horn is about rectangular in section,
being 0.25 oral diameter in length. Its free tip is squarely
truncated.

The thin, hyaline wall of the bowl is slightly thickened in
the collar.

Length, 26 to 36^1.

Acanthostomella obtusa differs from the other species in
having a stout, squarish aboral horn. The remaining species
have either no developed horn, or else a sharply pointed one.
Acanthostomella conicoides has fewer teeth (6 to 8) and is
more slender. Acanthostomella gracilis is alveolar, and has
a relatively longer, more cylindrical bowl, and no horn.

Recorded from two stations (45, 80) in the Galapagos
region, in a net and a pump sample taken at the surface and
at 50 meters, respectively. Frequency, 2 per cent at station 80.

52

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Temperature, 22?37-26?04 (24?2o); salinity, 35-23-35-94
(35.35); density, 23.75-24.30 (24.02); pH, 8.13-8.20 (8.16).

Petalotrichinae Kofoid and Campbell

Petalotrichineae, Kofoid and Campbell, 1929, p. 202.

Only the single genus Petalotricha belongs to this sub-
family.

PETALOTRICHA Kent emended

Petalotricha, Kofoid and Campbell, 1929, p. 202.

Large, smooth-walled, well finished loricae are character-
istic of Petalotricha. These are among the more complex
Tintinnoina of the warmest parts of the ocean. One species,
Petalotricha foli, is limited to the eastern Pacific, but P.
major is circumtropical.

Five species are described here.

Petalotricha ampulla (Fol) Kent
(Figure 26)

Petalotricha ampulla, Kofoid and Campbell, 1929, p. 203, fig.

389; Entz, 1935, pp. 15-26, figs. 1-15.
The stout lorica, with pointed aboral end, has a length of
1.06 oral diameters. The oral margin is irregularly serrate
with minute points. The upper collar is a horizontal shelf,
and the oral aperture has a diameter of but 0.87 the diameter
of the outer edge of this shelf. The lower collar is a basal
segment of a low inverted cone (28 ) with full sides; its
width is 0.22 oral diameter. Optically dark lines separate
the two collars and the bowl. The bowl rounds away from
the throat at 58 and at the shoulder reaches a diameter of
0.85 oral diameter. The bowl is 0.89 oral diameter at its
maximum, which lies at 0.4 oral diameter below the margin.
Below this level it contracts (at 32 for a distance of 0.52
oral diameter below the level of the maximum diameter, and
then at ioo° for the remaining distance). The aboral end is
broadly pointed but not prolonged.

The thin, uniform wall is not over 0.0 1 oral diameter in
thickness, and is grayish; there is a row of small oval lacunae
at the upper end of the lower collar. A double row of
lacunae also occurs below the shoulder.

The animal has about 100 small macronuclei, and 5 chro-
mosomes are reported in the micronuclei. If this last be
confirmed, the vegetative individual must be haploid, be-
cause the chromosome number is odd, and the reduction in
conjugation must be postzygotic, a condition unique in the
ciliates, although common in algae, mastigophorans, and
certain, if not all, sporozoans.
Length, no(.i.

Entz (1935) discusses the cytology of the animal in a
special paper.

Petalotricha ampulla is pointed aborally, unlike P. major,
which is rounded. Its bowl is short in contrast with that of
P. foli. in which the bowl is conical and elongated. It differs
from P. scrrata in its shallower, less regular, and less distinct
serrations and deeper throat. It lacks the curtain of P. entzi.
On the whole it is not likely to be confused with other com-
mon species, especially major and foli.

Recorded from eight stations, four each in the Atlantic
and the Pacific, as follows: two (4, 5) in the Atlantic drift,
two (14, 16) in the Gulf Stream, two ( 65, 66) in the South
Pacific middle latitudes, one (68) in the Galapagos region,
and one (113) in the North Pacific middle latitudes.

There are 1 pump and 8 net samples, of which 2 were
taken at 50 meters and 7 at 100 meters. Maximum fre-
quency, 30 per cent at station 68; other records above mini-
mum (2 to 10 per cent) from stations 4, 5, 14, 16, 65, 113;
averages in net samples, 5 and 12 per cent in the Atlantic
and Pacific, respectively.

Temperature: Atlantic, net samples i3?5i-23?64 (i5?77);
Pacific, pump sample 17^94, net samples i5?03-2i?74
(i7?77). Salinity: Atlantic, net samples 35.88-36.41 (35.95);
Pacific, pump sample 34.94, net samples 34.30-34.85 (34.60).
Density: Atlantic, net samples 24.84-27.01 (26.48); Pacific,
pump sample 25.26, net samples 24.06-25.52 (25.00). pH:
Atlantic, net samples 8.06-8.23 ( 8 -'4); Pacific, pump sample
8.12, net samples 8.10-8.23 (8.15).

Petalotricha capsa Brandt

Petalotricha capsa, Kofoid and Campbell, 1929, p. 203, fig. 387.

The saccular lorica, with flaring collars, goblet-shaped
bowl, and rounded aboral end, has a length of 0.93 oral
diameter. The oral margin is smooth, thin-edged, and
nearly erect. The upper and lower collars flare concur-
rently as basal segments of inverted truncated cones (62 ),
and each is laterally sigmoid. On the inside, the boundary
between the lower edge of the upper collar and the upper
edge of the lower collar is marked off by an erect, short
ledge, and there is a similar ledge at the lower edge of the
lower collar where it joins the bowl. On the outside, tiny
oval lacunae occur along the boundary lines of the two
collars. The diameter of the throat is 0.77 oral diameter.
The bowl below rounds over a slightly developed shoulder
and then becomes conical (10°) for about two-thirds the
length of the bowl, again increasing below this to 62 , and
finally to 125° near the aboral end. The aboral end is hemi-
spheroidal.

The wall is 0.02 oral diameter in thickness, and there are
thin, distinct inner and outer laminae between which are
enclosed two or three layers of rounded alveoles. These
alveoles are clearly visible externally, and the whole lorica
distinctly shows them. Large lacunae, of oval shape with the
axes directed vertically, occur in 2 or 3 rows in the upper
three-tenths of the bowl; they are 16 in number across one
face.

Length, 1251.1.

Petalotricha capsa resembles P. indica in having alveoles
in the wall in several layers. It differs from indica in its
lesser elongation, narrower aboral region, and more con-
stricted throat. From P. pacifica it may be distinguished by
its longer, less hemispherical bowl, and by the several layers
of alveoles. The remaining species have different wall struc-
ture.

Recorded from two stations (158, 159) in the region of
South Pacific island fields.

TAXONOMY AND DISTRIBUTION

53

There are 5 net samples, of which 1 was taken at the
surface, and 2 each at 50 and 100 meters. Maximum fre-
quency, 82 per cent at station 158 at 100 meters; average,
44 per cent.

Temperature. 27?8i-28?6o (28?i9); salinity, 35.58-35.89
( 55.74); density. 22.71-23.11 (22.90); pH, 8.37-8.39 (8.38).

Petalotricha foli Kofoid and Campbell

(Figure 27)
Petalotricka joli Kofoid and Campbell, 1929, p. 204, fig. 390.

The elongated conical lorica has a length of 2.34 oral
diameters. The oral margin is roughly serrate, with minute
points. The upper collar or shelf is a low basal segment of a
lull cone (90 ) with a width of less than 0.2 oral diameter
and with a lower diameter of nearly 0.87 oral diameter. The
lower collar is also a basal segment of a low, convex cone
(50 ) with a width of about 0.2 oral diameter and a lower
diameter of 0.77 oral diameter. There are sharp, optically
dark lines separating these two collars from each other and
from the bowl. The long bowl is a full cone (32 ). The
upper 0.16 of the bowl rounds with some suggestion of
shoulder which is about 0.82 oral diameter in diameter and
reaches 0.8 oral diameter near 0.2 total length below the
rim. Below this level the bowl contracts evenly, with some
minor local contractions and bulges, to the sharply pointed
but otherwise unmodified aboral end.

The wall is subuniformly about 0.02 oral diameter in
thickness, and has laminae and subrectangular prisms be-
tween the laminae. The lower end of the bowl (about 0.4
total length) has irregular vertical creases or folds, and near
the shoulder there are circular to narrow oval lacunae in two
or three rows. The remaining part of the lorica is almost
transparent, for the small prisms do not show an external
meshwork.

Length, 264 to 377H-

There are dwarf loricae of 1.75 oral diameters scattered
among the larger ones, and most of the loricae tend to be
stouter than the figured specimen (fig. 27).

Petalotricha joli is unlike the other species of the genus in
having a long, conical bowl, and in its heavy-set facies. Ap-
parently it spreads from off Mexico to the mid-Pacific.

Recorded from one station (151) in the North Pacific
trade region, in a net sample taken at 50 meters. Frequency,
11 per cent.

Temperature, i8?28; salinity, 34.42; density, 24.77; pH,
not recorded.

Petalotricha major Jorgensen
(Figures 24, 25)

Petalotricha major, Kofoid and Campbell, 1929, pp. 204-205,
fig. 384.

The rather short lorica has a globose bowl and a length of
only 0.9 oral diameter. The oral margin is serrate, with
minute points. The upper collar is a low, almost flat shelf
and flares ioo°. The oral aperture has a diameter of 0.86
that of the outer rim of the shelf. The lower collar is a basal

segment of an inverted truncated full cone (65 ) with a
diameter at its lower end of 0.7 oral diameter. The bowl is
globose. Its upper end is truncated where it joins the lower
collar, and it is 0.86 oral diameter in diameter at the maxi-
mum. The aboral end is smoothly rounded and without
differentiation.

The thin wall is uniformly about 0.01 oral diameter in
thickness, and there are thin laminae with a single layer of
rectangular prisms enclosed. The wall is glossy gray. There
is a single row of tiny oval lacunae at the upper end ol the
lower collar where it joins the upper collar, and a belt ol
several rows of scattered similar ones in the equatorial region
of the bowl.

Length, 95 to 1 iop.

An extreme form is figured (fig. 25) in which a minute
point is present. This lorica also has a very plump bowl
which is considerably shorter than wide. In the same col-
lection were other loricae much longer than wide. The com-
mon form in this collection conforms to type in every detail.
Some loricae have a suboral curtain comparable to that of
P. entzi.

Petalotricha major has a spherical bowl unlike that ol any
of the other species. It is quite unlike P. ampulla, in which
the aboral end is pointed, and it differs from P. entzi, which
also has a globose bowl, in that entzi has an outer hyaline
envelope developed around the collar. Petalotricha capsa has
a saccular bowl and alveoles in the wall in several layers.

There is a possibility that P. entzi belongs to this species,
since specimens with the diagnostic curtain occur. If this
should be proved, then major becomes a synonym of entzi,
since Kofoid's description of the latter was much earlier than
Jorgensen's of major.

Recorded from fifty-five stations, twenty-four in the At-
lantic and thirty-one in the Pacific, as follows: three (2, 14,
16) in the Gulf Stream, three (3, 4, 5) in the Atlantic drift,
six (17, 18, 19, 20, 20-21, 21) in the Sargasso Sea, eight (22,
23, 24, 25, 27, 28, 29, 30) in the Atlantic equatorial region,
four (31, 32, 33, 34) in the Caribbean Sea, three (46, 68, 69)
in the Galapagos region, four (63, 64, 65, 67) in the South
Pacific middle latitudes, eight (81, 82, 85, 94, 95, 96, 157.
160) in the region of South Pacific island fields, four (99,
152, 155, 156) in the Pacific equatorial region, three (100,

150, 151) in the North Pacific trade region, two (112, 113)
in the North Pacific middle latitudes, and seven (131, 135,
136, 137, 147, 148, 149) in the California region.

There are 15 pump and 58 net samples, of which 4 were
taken at the surface, 18 at 50 meters, 50 at 100 meters, and 1
at 150 meters. The preference of this species for subsurface
waters appears clear from the data. Maximum frequency,
86 per cent at station 30 at 100 meters; other records above
minimum (2 to 42 per cent) from stations 2, 3, 5, 16, 17, 18,
19, 20, 22, 23, 24, 27, 28, 31, 32, 33, 34, 46, 63, 65, 67, 68, 69,

82 > 8 5> 95. 9 6 > 99. I0 °- II2 - lr 3> '3 1 - *3 6 > W> r 4 8 > x 49> r 5°>

151, 152, 155, 156, 157, 160; averages in net samples, 16 and
12.3 per cent for the Atlantic and Pacific, respectively.

Temperature: Atlantic, pump samples i4?95~20?93 (i7?4o),
net samples i3?79-27?88 (2i?oo); Pacific, i5?03-28?74
(2^47) and n?48-28?74 (2i?8i), respectively. Salinity:

54

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Atlantic, pump samples 35.10-36.75 (35.82), net samples
35.88-37.05 (36.20); Pacific, 34.30-36.46 (35.01) and 33.36-
36.24 ( 35. u)* respectively. Density: Atlantic, pump samples
25.86-26.34 (26.09), net samples 23.26-27.01 (25.42); Pacific,
22.43-25.48 (24.29) and 22.31-26.50 (24.69), respectively.
pH: Atlantic, pump samples 7.96-8.20 (8.1 1), net samples
7.93-8.28 (8.23); Pacific, 8.08-8.34 (8.20) and 7.76-8.39
(8.21), respectively.

Petalotricha serrata Kofoid and Campbell
Petalolricha serrata Kofoid and Campbell, 1929, p. 205, fig. 386.

The lorica, with definitely serrated oral margin and a
saccular aborally pointed end, has a length of 1.1 oral diam-
eters. The oral margin has 48 short, sharp, erect, triangular
teeth. The upper collar flares (8o°) and its width is 0.09
oral diameter. Its lower edge is marked by a single row of
small, ovoid lacunae with horizontal axes. The lower collar
has a width of 0.12 oral diameter and is a band or cuff with
plane sides; it is differentiated only by being optically less
dense than the bowl. There is scarcely any nuchal con-
striction, and the bowl below reaches 0.9 oral diameter at a
similar distance below the margin. It contracts (26 ) below
the level of maximum diameter for a distance of two-thirds
its own length, and again more so (ioo°) below this to the
aboral end. The aboral end is distinctly pointed, with a
tiny nipple.

The gray wall is thin and has but a single layer of prisms
between the laminae. There are 17 large equatorial lacunae.

Length, 105 to 12011.

Petalotricha serrata resembles P. ampulla, but the oral
teeth are regular and pronounced. It has a cylindrical lower
collar, and unlike ampulla has scarcely any nuchal contrac-
tion. Its bowl is pointed, unlike that of P. major, and is
longer than that of P. paafica, but not so long as that of
P. indica. Petalotricha capsa has definite many-layered
alveolar walls, lacking in serrata.

Recorded from two stations, one each in the Atlantic and
the Pacific, as follows: one (30) in the Atlantic equatorial
region, and one (46) in the Galapagos region.

There are 2 net samples, one each from 50 and 100
meters. Frequency, minimum.

Temperature: Atlantic, 25754; Pacific, 23?26. Salinity:
Atlantic, 36.40; Pacific, 35.33. Density: Atlantic, 24.25;
Pacific, 24.13. pH: Atlantic, 8.28; Pacific, 8.16.

RHABDONELLIDAE Kofoid and Campbell
Rhabdonellidae Kofoid and Campbell, 1929, p. 206.

Included in this family are four genera: Protorhabdonclla ,
Epirhabdonclla, Rhabdonella. and Rhabdoncllopsis. Three
are found in the material of this expedition, and all are
tropical to subtropical.

PROTORHABDONELLA Jorgensen
Protorhabdonclla , Kofoid and Campbell, 1929, p. 206.

Protorhabdonclla is clearly the simplest genus of the Rhab-
donellidae in size and form. Kofoid and Campbell (1939,

p. 158) erected a new genus, Epirhabdonclla, with two new
species, also related but with finlike ribs.

Protorhabdonella is almost exclusively tropical and oceanic,
being most common in southern waters.

Three species are described here.

Protorhabdonella curta (Cleve) Jorgensen

Protorhabdonclla curta, Kofoid and Campbell, 1929, p. 207, fig.
39:5; Marshall, 1934, p. 646.

The tiny conical lorica, with numerous twisted ribs and
pointed aboral end, has a length of 1.8 oral diameters. The
oral margin is thin-edged and narrow, and an exceedingly
low cuff surrounds it. The conical bowl is 12 in the upper
half and increases to 68° in the lower part. There is a low,
rounded shoulder a little below the oral margin, the diameter
of which is 1.2 oral diameters; at its middle the bowl is
slightly less than 1.0 oral diameter. The upper bowl is
barely convex, but the lower is distinctly concave. The aboral
end is pointed, and conical (16 ) but not especially pro-
longed. There are about 24 strongly left-turned (15 ), very
low, linelike ribs. These ribs are rarely or never branched,
and die away some distance below the suboral shoulder;
they converge at the aboral end. Fenestrae are lacking.

The exceedingly thin wall is transparent.

Length, 39 to 52(1.

Protorhabdonella curta differs from P. simplex chiefly in
having more ribs that are strongly twisted, in contrast with
the fewer vertical ones of the other species. Differences in
shape are less marked, but some are of importance. Al-
though it has numerous ribs, it is not like Rhabdonella exilis,
R. indica, or R. amor, all forms with longer, narrower bowls
and emergent horns.

Recorded from four stations, one in the Atlantic and three
in the Pacific, as follows: one (31) in the Caribbean Sea, two
(54, 60) in the South Pacific middle latitudes, and one (113)
in the North Pacific middle latitudes.

There are 4 net samples, of which 1 was taken at the
surface and 3 at 100 meters. Frequency, 4 per cent at station
113, and 2 per cent at station 54; other records minimum;
average in the Pacific, 2.3 per cent.

Temperature: Atlantic, 22^56; Pacific, i4?97-2i?74( 1 8?48).
Salinity: Atlantic, 36.51; Pacific, 33.91-35.35 (34.64). Den-
sity: Atlantic, 25.22; Pacific, 24.06-25.37 (24.86). pH: At-
lantic, 8.19; Pacific, 8.07-8.23 (8.15).

Protorhabdonella simplex (Cleve) Jorgensen
(Figures 83, 84)
Protorhabdonella simplex, Kofoid and Campbell, 1929, p. 208,
fig. 395; Marshall, 1934, p. 646.

The short, conical lorica, with low, hyaline collar, few
bladelike fins, and pointed aboral end, has a length of 2.1S
oral diameters. The oral rim is simple, thin, and erect. The
low collar is a cuff with a length of little more than 0.01 oral
diameter. The conical bowl is about 9 within the upper 0.9
oral diameter, and increases to 42° in the lower 1.32 oral
diameters. Just below the collar is a barely evident rounded
shoulder, the diameter of which is 1.18 oral diameters; at the

TAXONOMY AND DISTRIBUTION

55

level at which the cone widens it is 1.04. At the aboral end
is a narrower (23 ), scarcely prolonged point which is the
homologue of the aboral horn of other species. There are
6 narrow (0.02 oral diameter), equidistant, decurrent blade-
like ribs which extend from the aboral end to the lower edge
of the collar. These ribs do not branch and are strictly
vertical.

The wall is hyaline and without fenestrae, and is thickest
just below the collar, where it reaches twice the width of the
cuff that rests on it; it gradually thins and is only three-tenths
as thick in the lower bowl. There are thin laminae, but
prisms were not evident between them.

Length, =52 to 98^1.

There is considerable variation in length and proportions,
probably correlated significantly with temperature. The
number of ribs (6 to 10) is interesting.

Protorhabdonella simplex resembles P. curta, but its ribs
are considerably fewer, and vertical. The bowl is longer and
narrows less quickly. It resembles P. ventricosa in having
few ribs, but that species is widely expanded toward the
aboral three-tenths (almost 1.4 oral diameters). It is unlike
Rhabdonella amor, in which the ribs are lower, twisted,
branched, and numerous, and which has a higher collar.

Recorded from eighteen stations, four in the Atlantic and
fourteen in the Pacific, as follows: two (20-21, 21) in the
Sargasso Sea, two (23, 24) in the Atlantic equatorial region,
six (41, 4=;, 46, 69, 77, 78) in the Galapagos region, four
(61, 63, 64, 66) in the South Pacific middle latitudes, two
(102, 109) in the North Pacific trade region, one (131) in
the California region, and one (143) in the North Pacific
middle latitudes.

There are 9 pump and 18 net samples, of which 10 were
taken at the surface, 7 at 50 meters, and 10 at 100 meters.
Maximum frequency, 8 per cent at station 61; other records
above minimum (2 to 5 per cent) from stations 21, 41, 45,
46, 69, 77, 78; averages in net samples, 1.3 and 3.1 per cent
for the Atlantic and Pacific, respectively; average in pump
samples, 1.5 loricae.

Temperature: Atlantic, net samples i5?55~26?57 (2i?88);
Pacific, pump samples i4?55~25?73 (20?72), net samples
i2?i2-23?72 (20?65). Salinity: Atlantic, net samples 35.61-
36.28 (36.03); Pacific, pump samples 34.39-36.17 (35.14),
net samples 33.36-36.04 (34.89). Density: Atlantic, net
samples 23.84-26.34 (24.98); Pacific, pump samples 23.65-
26.11 (24.63), net samples 24.06-25.31 (24.52). pH: At-
lantic, net samples 7.96-8.32 (8.17); Pacific, pump samples
7.92-8.30 (8.24), net samples 8.05-8.32 (8.15).

Protorhabdonella striatura Kofoid and Campbell
Protorhabdonella striatura Kofoid and Campbell, 1929, p. 20K,

fig- 392-
The elongated, conical, chalice-shaped lorica, with numer-
ous ribs and narrow conical aboral horn, has a length of
4.88 oral diameters. The oral margin is thin and erect. The
collar is a basal segment of a cone (21 °) and has a length of
about 0.5 oral diameter; its diameter at the lower end is
0.91 oral diameter. The elongated bowl gradually expands

from the throat to nearly 1.2 oral diameters near 1.65 oral
diameters below the rim. Below ibis level the bowl steadily
contracts (22 ) within less than 2.2 oral diameters, forming
an inverted segment of a cone. The pedicel is an inverted
asymmetrical cone (18 ) with a length of 1.46 oral diameters.

The rather transparent wall has 24 vertical ribs, a few of
which branch in the upper three-tenths of the bowl. There
are a few (6 to 8) small circular fenestrae scattered in a zone
within the posterior part of the bowl above the pedicel. The
wall is thin (about 0.02 oral diameter), with slight thicken-
ing at the throat.

Length, 125 to 165(1.

Protorhabdonella striatura differs from P. mira in having
a shorter pedicel and more ribs, and in contraction at the
throat. The other species of the genus have weaker devel-
opment of the pedicel.

Recorded from one station (27) in the Atlantic equatorial
region, in a pump sample taken at 50 meters. Frequency,
2 loricae.

Temperature, 26?04; salinity, 56.25; density, 23.98; pH,
8.30.

RHABDONELLA Brandt emended
Rhabdonclla, Kofoid and Campbell, 1929, pp. 209-211.

Rhabdonella is related to the much smaller, simpler Proto-
rhabdonella, and to the more complex Rhabdonellopsis. The
dominant structure in this genus is the vertical ribs. These
suggest the possible origin of the genus in some member of
the Favellidae or the Ptychocylidae, in which there are
many short surface pleats, and also suggest relationship to
the more advanced genera of the Epiplocylidae.

Rhabdonella occurs in tropical seas, there being little or
no limitation of species to any of the oceans save, perhaps, in
a few rare and little understood species, such as Rhabdonella
aberrans.

Eighteen species are described here.

Rhabdonella amor (Clcvc) Brandt
(Figure 85)

Rhabdonella amor, Kofoid and Campbell, 1929, p. 212, fig. 398;
Marshall, 1934, pp. 649-650, fig. i<>.

The small lorica has a high suboral rim, conical bowl, and
pointed aboral end; its length is 1.9 oral diameters. The oral
rim is ringlike, sharp, and high. The suboral trough is deep
and concave, and the diameter of its outer margin is 1.15 oral
diameters. There is little suboral flare. The bowl is dis-
tinctly conical (14 in the upper two-fifths, then 46°, and
finally 40 in the lowermost fifth). The aboral end is
pointed, sharp, and not prolonged.

The wall has a thickness of 0.1 oral diameter suborally,
and progressively thins in the bowl. There are thin laminae
which enclose several layers of exceedingly minute alveoles.
There are 32 to 44 left-deflected (as much as 12 ), commonly
branched and anastomosed ribs. The intercostae have 15 to
20 single or double rows of large oval fenestrae which pass
across the laminae and enclosed alveoles.
Length, 77 to 98a.

Marshall's 2 figured loricae are unusual in shape and in

56

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

faintness of fenestrae. They may, however, not be Rhab-
donella amor, but rather a new species peculiar to the Barrier
Reef.

Rlnil'donella amor averages fewer ribs than R. indica, is
longer, and lacks pronounced suboral Hare. It lacks the peg
of the somewhat longer R. cornucopia and the distinct horn
of the shorter R. exilis. It is easy to distinguish it from other
species.

Recorded from thirty-four stations, sixteen in the Atlantic
and eighteen in the Pacific, as follows: one (15) in the Gulf
Stream, two (19, 21) in the Sargasso Sea, nine (21-22, 22, 23,
24, 25, 27, 28, 29, 30) in the Atlantic equatorial region, four
(31, 32, 33, 34) in the Caribbean Sea, four (35, 35-36, 36, 39)
in the Pacific equatorial region, two (60, 62-63) in the South
Pacific middle latitudes, two (73, 80) in the Galapagos
region, six (81, 82, 85, 93, 96, 160) in the region of South
Pacific island fields, one (144) in the North Pacific middle
latitudes, one (146) in the California region, and two (150,
151) in the North Pacific trade region.

There are 19 pump and 41 net samples, of which 25 were
taken at the surface, 18 at 50 meters, and 17 at 100 meters.
In general this species has a pronounced surface preference.
Maximum frequency, 30 per cent at station 30; other records
above minimum (2 to 12 per cent) from stations 21-22, 22,
24, 25, 27, 29, 31, 32, 33, 34, 35, 35-36, 60, 80, 82, 85, 151, 160;
averages in net samples, 2.0 and 3.8 per cent for the Atlantic
and Pacific, respectively; the pump samples include from 1
to 15 loricae, with averages of 1.7 and 5.0 in the Atlantic and
Pacific, respectively.

Temperature: Atlantic, pump samples i4?6o-28?5i (23^59),
net samples i4?6o-27?88 (22^82); Pacific, i7? 4 6-2 9 ?3o
(24?39) and i4?97~28?58 (24^39), respectively. Salinity:
Atlantic, pump samples 35.22-36.99 (35.99), net samples
35.61-37.15 (36.29); Pacific, 34.59-35.42 (35-07) and 31.62-
36.33 (34.83), respectively. Density: Atlantic, pump samples
22.73-26.62 (23.77), net samples 23.26-26.34 (24.84); Pacific,
21.95-25. n (23.58) and 20.34-25.60 (23.33), respectively.
pH: Atlantic, pump samples 7.93-8.34 (8.31), net samples
7.93-8.31 (8.23); Pacific, 8.12-8.37 (8- 22 ) and 7.92-8.44
(8.23), respectively.

Rhabdonella brandti Kofoid and Campbell

(Figure 89)

Rhabdonella brandti Kofoid and Campbell, 1929, p. 213, fig.
400; Marshall, 1934, p. 649, fig. 24.

The short, chalice-shaped lorica, with tapering conical bowl
and fairly long horn, has a length of 3.42 oral diameters.
The oral rim is thin and erect, and its outer edge is the con-
cave upper margin of the inner face of the suboral trough.
The suboral trough is flattened toward the periphery, and
the diameter of its outer margin is 1.23 oral diameters. There
is some suboral flare (28°) within the anterior 0.3 oral diam-
eter. The bowl tapers (10°) for about 0.4 total length, then
becomes inverted subconical (33°) for approximately another
0.4. The aboral horn is narrow conical (13°), about 1 oral
diameter in length, and with a basal diameter of 0.23 oral
diameter. The free tip is sharply pointed.

The wall is thickest suborally (nearly 0.16 oral diameter)
and becomes gradually thinner lower down. There are thin
laminae which enclose minute alveoles. There are 48 long,
vertical, sometimes anastomosed ribs with 26 to 34 minute,
distinct fenestrae in each intercostal space. Minute alveoles
are externally evident in the generally brownish wall.

Length, 95 to 198(1.

This distinctive species has less suboral flare, less wide
bowl, and more irregular ribs than R. elegans. Its horn is
not so long or so narrow as that of R. quantula or R. inflata.

Recorded from four stations in the Atlantic, as follows:
two (15, 16) in the Gulf Stream, and two (19, 20) in the
Sargasso Sea.

There are 1 pump and 5 net samples, ot which 2 were
taken at the surface, 2 at 50 meters, and 2 at 100 meters.
Frequency, 2 per cent at station 19; 15 loricae in pump
sample at station 16; average for net samples, 1.6 per cent.

Temperature: pump sample 25^92, net samples 22^42-
24?8i (23?28). Salinity: pump sample 36.16, net samples
36.39-37.07 (36.73). Density: pump sample 23.93, net
samples 24.47-25.67 (25.17). pH: pump sample 8.24, net
samples 8.19-8.26 (8.22).

Rhabdonella conica Kofoiil and Campbell

Rhabdonella conica Kofoid and Campbell, 1029. pp. 214-215,
fig. 418.

The very tall, lanky lorica, with conical bowl, greatly
elongated horn, and little transition between bowl and horn,
has a length of 6.5 oral diameters. The oral rim is sub-
merged and recurved. The suboral trough is asymmetrically
concave; the diameter of its outer margin is 1.1 oral diam-
eters. The narrow bowl tapers (8°), but has a more or less
ringlike suboral swelling, its length being approximately 0.5
total length. The long aboral horn continues with the same
taper, although this is gradually reduced to less than 6° near
the tip.

The wall is only 0.03 oral diameter in thickness at the
thickest part. There is a suboral jelly-like curtain in some
individuals, in the anterior three-tenths or less. The laminae
are very thin and enclose extremely minute alveoles. There
are 48 left-deflected, often branched, and sometimes anasto-
mosed ribs. The intercostae have a dozen or more rather
large, oval, subequidistant fenestrae and a fine mesh work of
tiny hexagons.

Length, 290 to 480(1.

Some loricae reach 8.0 oral diameters in length, and the
aboral horn is often nearly transparent; mostly the loricae
are brown.

Because of its length and thinness, Rhabdonella conica is
easy to distinguish. In form it resembles R. aberrans, but
that species is shorter, with an irregular oral region and de-
fective ribs. Rhabdonella cuspidata is inflated in the lower
bowl and of stouter fades although about as long as conica.
The horn is always relatively longer in R. conica than in
R. spiralis, and there is almost no transition between bowl
and horn in conica; this character is rather distinct in spiralis.

Recorded from seventeen stations, one in the Atlantic and

TAXONOMY AND DISTRIBUTION

57

sixteen in the Pacific; it was also found off Easter Island.
The stations are as follows: one (19) in the Sargasso Sea,
three (47, 78, 80) in the Galapagos region, eight (48, 81, 82,
83, 84, 85, 95, 96) in the region of South Pacific island fields,
two (61, 65) in the South Pacific middle latitudes, and three
(107, 139, 140) in the North Pacific trade region.

There are 16 pump and 16 net samples, of which 9 were
taken at the surface, 16 at 50 meters, and 7 at 100 meters.
Maximum frequency, 31 per cent at station 78; other records
above minimum (2 to 14 per cent) from stations 47, 48, 61,
80, 81, 82, 85, 95, 139; average in Pacific net samples, 7.7
per cent.

Temperature: Atlantic, net sample 22?42; Pacific, pump
samples 23?88-29?34 (2f>?33), net samples io?92-28?74
(24^49). Salinity: Atlantic, net sample 37.05; Pacific, pump
samples 34.39-36.49 (35.74), net samples 34.05-36.49 (35.74).
Density: Atlantic, net sample 25.67; Pacific, pump samples
21.90-25. 11 (23.49), net samples 22.43-26.06 (23.91). pH:
Atlantic, net sample 8.25; Pacific, pump samples 8.14-8.39
(8.21), net samples 8.03-8.39 (8.21).

Rhabdonella cornucopia Kofoid and Campbell

Rhabdonella cornucopia Kofoid and Campbell, 1929, p. 215.
fig- 399-

The short lorica, with high oral rim, conical bowl, feeble
ribs, and conical aboral horn, has a length of 2.22 oral diam-
eters. The oral rim is erect, high, and ringlike, and the sub-
oral trough is flat; the diameter of the outer margin of the
latter is 1.22 oral diameters. There is almost no suboral
flare. The bowl is conical (18 increasing to 30 in the
lower part) and at its lower end gives rise to the aboral horn.
The horn is 0.55 oral diameter in length, conical (12 ),
truncated at the tip, and minutely open.

The wall is subuniformly 0.15 oral diameter in thickness.
There are thin laminae and enclosed faint alveoles. The
ribs number 26, and are very faint and difficult to follow, and
left-twisted (hardly io°); there are no fenestrae. The horn
is lemon yellow.

Length, 97 to 146(1.

Rhabdonella cornucopia is longer than R. cxilis, and has
faint ribs and a longer conical horn. Rhabdonella amor
lacks a horn and has distinct ribs. The remaining species
are altogether different.

Recorded from two stations in the Pacific, as follows: one
(60-61) in the South Pacific middle latitudes, and one (144)
in the North Pacific middle latitudes.

There are 1 pump and 1 net sample, both taken at the
surface. Frequency, minimum.

Temperature, net sample, 23^26; salinity, 34.97; density,
23.86; pH, 8.37; all records at station 144.

Rhabdonella cuspidata (Zacharias) Brandt

Rhabdonella cuspidata, Kofoid and Campbell, 1929, p. 215, rig.
4'7-

The greatly elongated lorica, with contraction and later
inflation and conical horn, has a length of 7.0 oral diameters.
The oral rim is low, ringlike, and submerged by the higher

outer margin of the concave suboral trough; the diameter of
this margin is 1. 17 oral diameters. The bowl contracts (10°)
for 1.33 oral diameters, basing .11 the lower end a diameter
of 0.9 oral diameter, and then expands, reaching 1.0 oral
diameter at 0.46 total length below the rim. The lower bowl
is convex conical (30 ) within [.5 oral diameters. At its
lower end is the narrow conical (8 ) aboral horn, the
length ol which is nearly 0.36 total length; its free tip is
sharp.

The wall has a thickness of 0.1 oral diameter across the
suboral trough and becomes reduced to a third of that in
the bowl. There are thin laminae which enclose several
layers of small alveoles, and the fenestrae cut across from
lamina to lamina. There are approximately 30 subvertical,
sometimes branched ribs. The intercostae have 12 to 30
very minute, oval, faint fenestrae.

Length, 421U.

Many loricae are much shorter (down to 250(1) than
usual.

Rhabdonella cuspidata, with its lateral concavity and swell-
ing and its great length, is easily distinguished from other
species. In elongation and proportions it approaches R.
conica, but it is otherwise only remotely like that species.

Recorded from one station (65) in the South Pacific middle
latitudes, in a net sample taken at the surface. Frequency,
8 per cent.

Temperature, 2o?22; salinity, 34.53; density, 24.37; pH,
8.10.

Rhabdonella elegans Jorgensen emended Kofoid
and Campbell

Rhabdonella elegans, Kofoid and Campbell, 1929, p. 215, fig.
401.

The rather short lorica, with wide, generally conical bowl
and tapering horn, has a length of 2.81 oral diameters. The
oral rim is relatively high, with outward-sloping sides. The
suboral trough is deep and subangular; the diameter of its
outer margin is 1.17 oral diameters. There is a distinct flare
(62°) well within the anterior 0.5 oral diameter. The
diameter of the bowl at the lower end of the flare is 1.0 oral
diameter. Below this level the bowl is decidedly convex
conical (io° increasing to 42 in the lower half). The aboral
horn is narrow conical (12 ), and 0.82 oral diameter in
length; it has a sharply pointed free tip.

The wall has a thickness of 0.06 oral diameter in the bowl
and has very little suboral thickening. There are thin
laminae which enclose fine primary alveoles in several layers.
There are approximately 24 vertical, unbranched ribs run-
ning from end to end. Fenestrae are lacking.

Length, no to 122U.

Rhabdonella elegans differs from R. cornucopia in its
longer horn and more convex bowl. It has a horn, which is
lacking in R. amor, and the horn is longer than that of R.
indica. It is much longer and more convex than R. exilis.

Recorded from one station (14) in the Gulf Stream, in a
net sample taken at 100 meters. Frequency, minimum.

Temperature, i4?02; salinity, 35.59; density, 26.66; pH,
8.06.

5«

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

dency toward the formation of a spindle near the distal end,

Rhabdonella exilis Kofoid and Campbell

Rhabdonella exilis Kofoid and Campbell, 1929, p. 216, fig. 396.

The very short lorica, with high oral rim, narrow conical
bowl, and pointed, prolonged peg-shaped horn, has a length
of 2.52 oral diameters. The oral rim is high and ringlike;
the suboral trough is plane, and angled out to the low outer
margin of the trough, the diameter of which is 1.09 oral
diameters. There is no suboral flare. The conical bowl
(io° to 15 in the upper part, increasing to as much as 45°
in the lower part) contracts evenly to the aboral end. The
aboral horn is nearly 0.5 oral diameter in length, peg-shaped,
and distally pointed.

The wall is thickest suborally, gradually thinning in the
lower bowl. Laminae are evident only in the upper bowl,
those of the lower bowl being apparently fused. There are
46 left-directed (18°), weak, unbranched ribs which con-
tinue from end to end. Fenestrae are lacking.
Length, 60 to jo\i.

Rhabdonella exilis is shorter than R. cornucopia, and has
more ribs and also stouter proportions. The horn is better
developed than in R. indica.

Recorded from twelve stations, three in the Atlantic and
nine in the Pacific, as follows: one (20) in the Sargasso Sea,
two (33, 34) in the Caribbean Sea, one (36) in the Pacific
equatorial region, three (63, 64, 65) in the South Pacific
middle latitudes, one (78) in the Galapagos region, two (91,
159) in the region of South Pacific island fields, one (140)
in the North Pacific trade region, and one (146) in the Cali-
fornia region.

There are 10 pump and 3 net samples, of which 6 were
taken at the surface, 5 at 50 meters, and 2 at 100 meters.
Frequency, 3 per cent at station 65; 2 per cent at stations 36,
140, 146; average for Atlantic net samples, 1.6 per cent.

Temperature: Atlantic, pumpsamples26?05-28?5i (27^28),
net samples 23?i7-2 4 ?g8 (2 4 ?o7); Pacific, i5?90-28?65
(22^64) and 28?6o, respectively. Salinity: Atlantic, pump
samples 35.87-36.55 (36.21), net samples 36.49-36.53 (36.51);
Pacific, 31.62-36.03 (34.54) and 35.74, respectively. Density:
Atlantic, pump samples 22.91-24.21 (23.56), net samples
24.52-25.03 (24.77); Pacific, 20.34-25.43 (23.60) and 22.77,
respectively. pH: Atlantic, pump samples 8.28-8.37 (8-3 2 )>
net samples 8.18-8.21 (8.19); Pacific, 8.10-8.39 (8.19) and
8.37, respectively.

Rhabdonella hebe (Cleve) Brandt

Rhabdonella hebe, Kofoid and Campbell, 1929, p. 216, fig. 409.

The moderately tall lorica, with moderate number of ribs,
conical bowl, and spindle-like swelling on the horn, has a
length of 4.15 oral diameters. The oral rim is very low, erect,
and ringlike, and the suboral trough which surrounds it is
concave; the outer margin has a diameter of 1.26 oral diam-
eters. The suboral flare (58 ) is within 0.1 oral diameter
and forms a thickened suboral lip around the upper end of
the conical bowl. The bowl is subcorneal (20 in the anterior
0.37 total length and 36 in the lower 0.19). The aboral
horn is conical (8°), 0.44 total length in length, with a ten-

and with a minute, truncated, open ( ? ) free tip.

The wall is thickest across the suboral shelf, where it
reaches nearly 0.16 oral diameter; it becomes less than three-
tenths as much in the bowl. There are thin laminae which
enclose several layers of small, hexagonal prisms. Fenestrae
connect the laminae across the wall at intervals. There is a
curtain of jelly-like material enclosing the upper 0.24 of the
lorica. There are 1 1 to 28 left-deflected (5 to 7 ), branched
ribs which die away on the suboral flare. Fenestrae are scat-
tered in 1 or 2 rows up and down each intercosta. The horn
is denser than the nearly transparent bowl.

Length, 200 to 330^.

Rhabdonella hebe may be at once distinguished from R.
spiralis by shorter length, fewer ribs, proportions, and bowl
shape. Its horn is more distinctly set off from the bowl than
in R. valdestriata; it has fenestrae, lacking in R. chavesi, and
its oral region is narrower than in R. striata. It is not likely
to be confused with other species.

Recorded from seven stations, three in the Atlantic and
four in the Pacific, as follows: one (15) in the Gulf Stream,
one (30) in the Atlantic equatorial region, one (31) in the
Caribbean Sea, one (45) in the Galapagos region, one (57)
in the South Pacific middle latitudes, one (145) in the North
Pacific middle latitudes, and one (146) in the California
region.

There are 6 pump and 4 net samples, of which 3 were
taken at the surface, 3 at 50 meters, and 4 at 100 meters.
Maximum frequency, 72 per cent at station 15; other records
above minimum (6 to 12 per cent) from stations 145, 146 in
pump samples; average in Pacific pump samples, 24.6 loricae.

Temperature: Atlantic, pump samples 1 8?56-i9?27(i8?9i),
net samples 22? 5 6-2 7 ?88 (2 5 ?o8); Pacific, i6? 5 8-22? 3 7
(20?05) and 22?37, respectively. Salinity: Atlantic, pump
samples 36.08-36.51 (36.32), net samples 36.45-36.47 (36.46);
Pacific, 34.32-34.91 (34.61) and 35.23, respectively. Den-
sity: Atlantic, pump samples 26.12-26.26 (26.19), net samples
23.26-25.22 (24.31); Pacific, 23.88-24.66 (24.37) and 24.30,
respectively. pH: Atlantic, pump samples 8.19-8.21 (8.20),
net samples 8.19-8.30 (8.25); Pacific, 8.14-8.31 (8.26) and
8.13, respectively.

Rhabdonella henseni ( Brandt) Brandt
(Figure 90)

Rhabdonella henseni, Kofoid and Campbell, 1929, p. 216, fig.
408.

The moderately elongated lorica, with wide, chalice -shaped
bowl, rather long, thick aboral horn, and vertical ribs, has 1
length of 5.1 oral diameters. The oral rim is hyaline, higher
than the margin of the suboral trough, thin, ringlike, and
erect. The suboral trough is shallow, and its outer margin
has a diameter of 1.13 oral diameters. There is scarcely any
suboral flare. The bowl is subcylindrical for nearly 0.4(1 total
length, and convex conical (40 ) for about 1.1 oral diam-
eters. The elongated, conical (8°) aboral horn has a basal
diameter of 0.3 oral diameter, and a length of almost 0.33
total length; its junction with the bowl is often asymmetrical.

TAXONOMY AND DISTRIBUTION

59

The wall is thickest across the ridgelike suboral Hare,
where it reaches 0.05 oral diameter; elsewhere it is decidedly
thinner. There are exceedingly thin, dark laminae with
numerous minute alveoles enclosed. A curtain-like film of
transparent jelly encloses the upper bowl. There are 18
vertical, frequently anastomosing, branching, rarely discon-
tinuous and short ribs, which run the whole length ot the
lorica save suborally. There are 12 to 18 small fenestrae (the
sites of coccoliths), and fine alveoles form an external mesh-
work in the intercostae. The whole lorica is a deep brown
color.

Length, 303ft.

The aboral horn of the figured specimen (fig. 90) is not
so thick as usual.

Rhabdonella henseni is stouter and plumper than any
species of the R. spiralis group. It is closest to R. torta and
R. lohmanni. It may be distinguished from lohmanni by the
vertical rather than oblique ribs, the smaller size, and the
more slender horn. From torta it may be distinguished by
the larger size and the vertical rather than right-spiral ribs.

Recorded from three stations, one in the Atlantic and two
in the Pacific, as follows: one (17) in the Sargasso Sea, one
(96) in the region of South Pacific island fields, and one
(135) in the California region.

There are 2 pump samples and 1 net sample, of which 2
were taken at the surface and 1 at 50 meters. Frequency,
minimum.

Temperature: Atlantic, net sample 2i?85; Pacific, pump
samples 23?70-29?30 (26?53). Salinity: Atlantic, net sample
36.60; Pacific, pump samples 35.12-35.27 (35.19). Density:
Atlantic, net sample 25.49; Pacific, pump samples 22.19-23.83
(23.01). pH: Atlantic, net sample 8.27; Pacific, pump sam-
ples 8.23-8.37 (8.30).

Rhabdonella indica Laackmann
Rhabdonella indica, Kofoid and Campbell, 1929, p. 217, fig. 397.

The short lorica, with conical bowl, short, peglike aboral
horn, and numerous ribs, has a length of 2.15 oral diameters.
The oral rim is high, erect, ringlike, and sharp-edged. The
suboral trough is concave; the diameter of its outer margin
is 1. 17 oral diameters. There is a distinct suboral flare
(35°) within the upper 0.1 oral diameter. The bowl is
convex conical (15° in the anterior half, increasing to 45 in
the posterior section, and then 20 ). The aboral end has a
prolonged, peglike horn with distal sharp point.

The wall reaches 0.09 oral diameter in thickness suborally,
but thins rapidly in the middle. There are thin laminae
which enclose several layers of minute alveoles. There are
36 to 42 delicate, left-twisted (15°), sometimes branched
ribs, which die away on the swollen suboral flare. Fenestrae
are lacking.

Length, 56 to 631.1.

Rhabdonella indica differs from R. amor in size, lack of
fenestrae, twist of the ribs, and more numerous ribs. It is
somewhat shorter than R. exilis, with fewer ribs and greater
density. It can scarcely be confused with other species.

Recorded from twenty stations in the Pacific, as follows:

three (36, 37, 39) in the Pacific equatorial region, four (46,
47, 78, 80) in the Galapagos region, three (48, 82, 90) in the
region of South Pacific island fields, one (58) in the South
Pacific middle latitudes, three ( 107, 109, 150) in the North
Pacific trade region, four (135, 1 ?C>, 146, 147) in the Cali-
fornia region, and two (142, 145) in the North Pacific
middle latitudes.

There are 24 pump and 6 net samples, of which 16 were
taken at the surface, 9 at 50 meters, and 5 at 100 meters. Max-
imum frequency, 6 per cent at station 46; other records above
minimum (2 to 5 per cent) from stations 48, 78; average in
net samples, 42 per cent; 2 to 15 loricae in pump samples.

Temperature: pump samples I4?40-27?I2 (23^27), net
samples i6?98-24?38 (22?57). Salinity: pump samples 31.62-
36.44 (35.34), net samples 33.97-36.44 (35.50). Density:
pump samples 20.20-26.07 (23.35), net samples 24.11-24.86
(24.45). pH: pump samples 7.85-8.32 (8.28), net samples
8.12-8.23 ( 8 -'7)-

Rhabdonella inflata Kofoid and Campbell

Rhabdonella inflata Kofoid and Campbell, 1929, p. 217, fig. 403.

The short, rather stout lorica, with chalice-shaped bowl
and wide aboral region, with little transition, has a length of
3.66 oral diameters. The oral rim is low, ringlike, and erect.
The suboral trough is flat and the diameter of its outer mar-
gin is 1.22 oral diameters. There is very little suboral flare.
The bowl tapers (5 ) for nearly 0.4 total length and then
becomes subcorneal (33°) within 1.2 oral diameters. The
transition between lower bowl and horn is gradual. The
aboral horn has a length equal to that of the lower bowl, is
subconical (14°), and is sharply pointed at the free tip.

The wall thickness reaches not over 0.06 oral diameter
suborally and thins down in the bowl. There are thin
laminae which enclose small alveoles in several layers. There
are upwards of 60 subvertical, mostly unbranched ribs, and
the intercostae have many small, faint fenestrae.

Length, 124 to 20op.

Rhabdonella inflata is shorter than R. spiralis, with grad-
ually differentiated horn and with more ribs. In similar
characters it differs from R. hebe, which last has a spindle-
like knob lacking in inflata. It is stouter than R. striata and
has fainter fenestrae.

Recorded from two stations (131, 147) in the California
region, in net samples taken at 100 meters. Frequency, 2
per cent at station 131.

Temperature, I2?i2-i9?27 (i5?69); salinity, 33.36-35.04
(34.20); density, 25.00-25.31 (25.15); pH, 8.29-8.32 (8.30).

Rhabdonella lohmanni Kofoid and Campbell

Rhabdonella lohmanni Kofoid and Campbell, 1929, p. 218, fig.
416.

The tall lorica, with chalice-shaped bowl, oblique ribs, and
stout horn, has a length of 5.5 oral diameters. The oral rim
is low, ringlike, and slanted outward. There is almost no
suboral trough or suboral flare; the diameter of the bowl at
the upper end is, however, 1.11 oral diameters. The upper
two-thirds of the bowl is subcylindrical, and the lower bowl

6o

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

is subcorneal (up to 35°). The aboral horn reaches 0.38 total
length and is relatively stout subconical (9°). Its free tip is
pointed.

The wall reaches a maximum thickness of only about 0.05
oral diameter. There are thin laminae which enclose small
alveoles. There are 48 definitely right-turning (io°),
branched, often anastomosed, heavy ribs, which fade away
near the distal tip of the horn and just below the suboral
rim; there are no fenestrae.

Length, 317 to 37711.

Rlmbdonella lohmanni resembles R. henseni in general, but
has oblique ribs and a stout horn, and is longer. Rhabdonella
torta is shorter (210 to 22811) but otherwise rather similar.

Recorded from two stations (138, 140) in the North Pacific
trade region, in 2 pump samples, 1 taken at the surface and
1 at 50 meters. Frequency, minimum.

Temperature, 26?i4~26?87 (26750); salinity, 34.85-35.02
(34.93); density, 22.80-22.90 (22.85); P H , 8.35-8.39 (8.37).

Rhabdonella poculum (Ostenfeld and Schmidt) Brandt
(Figure 92)

Rhabdonella poculum, Kofoid and Campbell, 1929. p. 218, fig.
405.

The short, wide lorica, with nearly cylindrical upper and
conical lower bowl, and wide conical horn, has a length of
1.93 oral diameters. The oral rim is entire, and is a cufl
which rises above the suboral trough which surrounds it.
The suboral trough is shallow and concave; the diameter of
its outer margin is 1.1 oral diameters. The bowl flares (10 )
within the anterior 0.1 oral diameter, then becomes cylin-
drical for 0.85 oral diameter, and finally convex conical
(6o°) for nearly 0.54 oral diameter. The aboral horn is in-
verted and concave conical (40 ), with a basal diameter of
0.32 oral diameter, and with a sharply pointed free tip.

The wall is subuniformly about 0.09 oral diameter in
thickness. There are 24 deflected (15 ) ribs, which arise on
the horn or near it, and die away just below the suboral
flare. The ribs are often branched and sometimes anasto-
mosed, and the intercostal regions have 8 to 12 scattered,
oval fenestrae. The lorica is commonly dark brown.

Length, 85"..

The Carnegie loricae show much more transition between
lower bowl and horn than is shown in the single figure of
Kofoid and Campbell; they have fewer ribs (left, rather than
right, deflection), and have fenestrae.

Rhabdonella poculum may be distinguished at once from
other species by the general form. In some ways it is like
R. amor, but it has a distinctly conical, well differentiated
horn. The shortness, and the basal width of the horn arc-
characters which are especially distinct.

Recorded from three stations in the Pacific, as follows:
two (40, 41) in the Galapagos region, and one (93) in the
region of South Pacific island fields.

There are 3 pump and 3 net samples, of which 4 were
taken at the surface, 1 at 50 meters, and 1 at 100 meters.
Maximum frequency, 13 per cent at station 41; average in
net samples, 5.3 per cent.

Temperature: pump samples 20^42-28^74 (24745), net
samples 14755-20742 (16776). Salinity: pump samples 33.70-
34.71 (34.20), net samples 34.19-35.02 (34.40). Density:
pump samples 21.95-24.06 (23.07), net samples 24.06-26.11
(25.33). pH: pump samples 8.1 1-8.30 (8.14), net samples
7.87-8. 1 1 (7.96).

Rhabdonella quantula Kofoid and Campbell

Rhabdonella quantula Kofoid and Campbell, 1929, p. 218, fig.
402.

The rather short lorica, with narrow, chalice-like bowl and
elongated horn, has a length of 3.66 oral diameters. The
oral rim is low and ringlike, and extends only slightly above
the suboral trough. The suboral trough is concave and the
diameter of its outer margin is 1.22 oral diameters. There is
some suboral flare (35°) within the upper 0.05 oral diameter.
The bowl tapers (16 ) in the anterior 0.37 total length and
then becomes subconical (30 ) within the posterior 0.35
total length. The aboral horn (0.29 total length in length)
is narrow conical (5 ) and sharply pointed at the free tip.

The wall is thickest across the suboral flare, where it
reaches 0.1 oral diameter; at other levels it is reduced by
one-half. There are thin laminae which enclose fine alveoles.
There are upwards of 54 subvertical, continuous, distinct,
unbranched ribs, and in each intercostal area are 6 to 8 small
circular fenestrae.

Length, 138 to 172U.

Rhabdonella quantula differs from R. spiralis mainly in
shortness and in greater number of ribs. Rhabdonella inflata
is of about the same size, but has a longer, less tapering
bowl; R. cornucopia has a stubby aboral horn. The lower
bowl of R. brandti is wider, and its horn is shorter than in
quantula.

Recorded from ten stations in the Pacific, as follows: three
(35, 35-36, 99) in the Pacific equatorial region, two (78, 80)
in the Galapagos region, four (81, 82, 84, 85) in the region
of South Pacific island fields, and one (140) in the North
Pacific trade region.

There are 3 pump and 10 net samples, of which 3 were
taken at the surface, 7 at 50 meters, and 3 at 100 meters.
Maximum frequency, 75 per cent at station 35-36; other
records above minimum (2 to 8 per cent) from stations 35,
82, 85, 99; average in net samples, 10.6 per cent.

Temperature: pump samples 25794-28732 (26777). net
samples 14733-27789 (23748). Salinity: pump samples
35.02-35.95 (35.58), net samples 34.88-36.42 (35.71). Den-
sity: pump samples 22.45-23.75 (23.09), net samples 22.50-
26.06 (24.09). pH: pump samples 8.16-8.34 (8.23), net
samples 7.88-8.22 (8.13).

Rhabdonella spiralis (Fol) Brandt emended Kofoid
and Campbell

Rhabdonella spiralis, Kofoid and Campbell, 1929, p. 219, fig.
414; Hofker (part), 1931, pp. 378-381, figs. 68-72 (for fig.
67 see Rhabdonellopsis triton); Marshall, T934, pp. 646-648,
figs. 23, 27(F).

The moderately tall, elongated lorica, with tapering bowl

TAXONOMY AND DISTRIBUTION

61

and horn, subequal in length, has a length of 5.5 oral diam-
eters. The oral rim is erect, ringlike, and about as high as
the outer rim of the suboral trough. The suboral trough is
concave and the diameter of its outer margin is 1.2 oral
diameters. The bowl flares (38 ) within the upper 0.1 oral
diameter. It then tapers (14°) for 0.42 total length, then
becomes subcorneal (28 ) for 0.18. From its lower end
arises the aboral horn. The horn tapers (7 ) and has a
length of 0.4 total length, and its free tip is sharp.

The wall has a thickness of not over 0.1 oral diameter
across the trough, and thins regularly below. There are thin
laminae which enclose several layers of minute alveoles.
There are 42 sometimes branched, anastomosed, subvertical
ribs, which continue from end to end. Between the ribs are
12 to 20 small, scattered circular fenestrae, mostly gathered
in the upper bowl and more rare below.
Length, 260(1.

That part of Hofker's material which has distal knobs is
assigned in this report to Rhabdonellopsis niton. Marshall
does not distinguished spiralis from related species, but since
only one figure is clearly of this species, this Barrier Reef
material is all included as spiralis. One of Marshall's figures
(fig. 27) strongly suggests Rhabdonella anadyomene , save
that the ribs are reversed and the oral rim more developed;
this lorica may, however, be only an incomplete (abnormal
or defective?) specimen. It is not at all like the imperfect
R. hydria, which is probably only a defective spiralis. Defec-
tive loricae were not encountered in the Carnegie material
save at station 1, where at 70 meters one specimen somewhat
like Marshall's was found. On the whole it is rather re-
markable that so few defective loricae of these ciliates are
found, considering their abundance in the ocean. Perhaps
this is because they are quickly formed.

Rhabdonella spiralis is closest to R. valdestriata, R. striata,
and R. hebe. It is usually longer than these, with more
slender proportions, with distinct suboral flare, and with
subequal upper bowl and horn. There is more transition
between lower bowl and horn than in hebe or valdestriata,
but not so much as in striata. It never has the length, nar-
rowness, or facies of R. conica. The remaining species are
clearly unlike it and occasion no opportunity for confusion.
Recorded from forty-four stations, sixteen in the Atlantic
and twenty-eight in the Pacific, as follows: three (1, 15, 16)
in the Gulf Stream, four (17, 18, 19, 21) in the Sargasso Sea,
five (24, 26, 27, 29, 30) in the Atlantic equatorial region,
four (31, 32, 33, 34) in the Caribbean Sea, two (35, 37) in
the Pacific equatorial region, eight (45, 46, 47, 68, 69, 78, 79,
80) in the Galapagos region, six (48, 87, 89, 90, 95, 97) in
the region of South Pacific island fields, nine (50, 51, 53, 54,
61, 62, 62-63, 64, 65) in the South Pacific middle latitudes,
two (131, 146) in the California region, and one (145) in
the North Pacific middle latitudes.

There are 46 pump and 27 net samples, of which 31 were
taken at the surface, 24 at 50 meters, 1 at 70 meters, and 17
at 100 meters. The preference of this species for surface
water is evident. Maximum frequency, 90 per cent at station
64; other records above minimum (2 to 88 per cent) from
stations 18, 30, 31, 54, 61, 62-63, °5> : 45! average in net

samples, Atlantic 1.6 per cent. Pacific 37.S per cent; in pump
samples, 300 loricae counted at station 15; other pump
records 1 to 43, average in the Pacific 6.0.

Temperature: Atlantic, pumpsamplcs i8?40-28?5i (25?76),
net samples i5? 55 -28?o 5 (22^28); Pacific, i3?28-28? 74
(22^70) and io?92-24?38 (i8?5o), respectively. Salinity:
Atlantic, pump samples 29.70-36. 58 (35.80), net samples
35.61-37.15 (36.45); Pacific, 31.68-36.44 (35.27) and 33.36-
36.03 (34.59), respectively. Density: Atlantic, pump samples
18.62-26.01 (23.69), net samples 23.20-26.34 (25.00); Pacific,
20.20-25.40 (24.15) and 24.24-26.06 (24.58), respectively.
pH: Atlantic, pump samples 8.11-8.31 (*.z(*). net samples
7.96-8.37 (8.29); Pacific, 8.05-8.28 (8.17) and 8.03-8.34
(8.17), respectively.

Rhabdonella striata (Biedermann) Brandt emended Kofoid
and Campbell

Rhabdonella striata, Kofoid and Campbell, 1929, p. 219, fig.

411.
The moderately tall lorica, with stout conical bowl and
elongate conical horn, has a length of 3.71 oral diameters.
The oral rim is a low, outward-recurved rim, which projects
a little above the outer margin of the suboral trough. The
suboral trough is flattened concave; the diameter of its outer
margin is 1.09 oral diameters. There is a wide suboral flare
(6o°) within the upper 0.1 oral diameter. The upper bowl
tapers (16 ) for 0.42 total length, and then becomes sub-
conical (44°) for 0.19 total length. The aboral horn is
narrow concave conical (5 ), with a length of 0.39 total
length and with a truncated free tip.

The wall has a maximum thickness of approximately 0.1
oral diameter. There are thin laminae which enclose fine
alveoles in several layers. There are 42 commonly branched,
anastomosed, subvertical, low ribs which continue from end
to end. Fenestrae are numerous, but small and not espe-
cially conspicuous.
Length, 225(1.

Rhabdonella striata has a more sharply differentiated lower
bowl and more ribs than R. valdestriata. It is stouter and
shorter, and has more ribs than R. spiralis. Rhabdonella
brandti is shorter.

Recorded from fifty-five stations, sixteen in the Atlantic
and thirty-nine in the Pacific, as follows: three (20, 20-21,
21) in the Sargasso Sea, nine (22, 23, 24, 25, 26, 27, 28, 29,
30) in the Atlantic equatorial region, four (31, 32, 33, 34) in
the Caribbean Sea, six (52, 56, 63, 65, 66, 67) in the South
Pacific middle latitudes, three (68, 69, 80) in the Galapagos
region, seven (89, 90, 93, 95, 96, 97, 159) in the region of
South Pacific island fields, four (99, 152, 153, 154) in the
Pacific equatorial region, six (100, 107, 108, 109, no, 138)
in the North Pacific trade region, eight (132, 133, 135, 136,
137, 146, 148, 149) in the California region, and five (141,
142, 143, 144, 145) in the North Pacific middle latitudes.
Rhabdonella striata is the most common Rhabdonella in the
ocean and is most widely distributed in warmer seas.

There are 54 pump and 45 net samples, of which 37 were
taken at the surface, 24 at 50 meters, and 38 at 100 meters.

62

OCEANIC T1NTINNOINA OF LAST CRUISE OF CARNEGIE

Maximum frequency, 78 per cent at station 20-21; other
records above minimum (2 to 60 per cent) from stations 21,
22, 23, 24, 25, 27, 28, 29, 30, 32, 33, 34, 68, 69, 80, 99, 136,
137, 145, 152, 153, 154, 159; averages in net samples, 17.9
and 4.8 per cent for the Atlantic and Pacific, respectively; in
pump samples, 150 loricae at station 96; average in Pacific
pump samples, 10.3.

Temperature: Atlantic, pump samples i4?6o-27?6i (23? 1 1 ),
net samples i4?6o-28?5 4 (23?54); Pacific, i5?84-29?43
(23?68) and n?48-28?6o (ai?47), respectively. Salinity:
Atlantic, pump samples 35.22-36.75 (35.96), net samples
35.61-36.65 (36.13); Pacific, 33.68-35.39 (35.08) and 34.30-
35.95 (34.96), respectively. Density: Atlantic, pump samples
22.84-26.62 (24.43), net samples 21.78-26.02 (23.49); Pacific,
21.70-25.48 (23.45) and 22.31-26.50 (24.31), respectively.
pH: Atlantic, pump samples 7.93-8.32 (8.20), net samples
7.93-8.32 (8.19); Pacific, 8.08-8.47 (8.29) and 7.76-8.39
(8.16), respectively.

Rhabdonella torta Kofoid and Campbell
Rhabdonella torta Kotoid and Campbell, 1929, p. 220, fig. 404.

The fairly long lorica, with stout, chalice-shaped bowl,
oblique ribs, and wide conical horn, has a length of 4.0 oral
diameters. The oral rim is submerged and low. The sub-
oral trough is flat and the diameter of its outer rim is 1.18
oral diameters. The bowl tapers (15°) in the anterior two-
thirds and becomes subconical (30 ) in the lower section.
The aboral horn is conical (22 ), little differentiated, and
about 0.3 total length in length, and has a blunted, free tip.

The wall reaches 0.11 oral diameter in thickness suborally
and thins regularly in the bowl. There are thin laminae and
enclosed alveoles in several layers. There are 56 right-twisted
(upwards of 40 ), feebly anastomosed and branched ribs;
the tips of the ribs are left-turned near the suboral margin.
There are 4 to 8 distinct fenestrae in the intercostae. The
lorica, as in Rhabdonella lohmanni and R. hensem, is deep
brown.

Length, 210 to 228U..

Rhabdonella torta is related to R. henseni and R. lohmanni,
but is much shorter than either and has a much less clearly
differentiated horn. The ribs are also different.

Recorded from station 61 in the South Pacific middle lati-
tudes, in a net sample taken at the surface. Frequency, 4
per cent.

Temperature, i6?9o; salinity, 34.05; density, 24.83; pH,
8.05.

Rhabdonella valdestriata Brandt

(Figure 91)

Rhabdonella valdestriata. Kofoid and Campbell, 1929, p. 220,
fig. 410.

The moderately tall, chalice-shaped lorica, with almost no
suboral flare, slight transition between bowl and horn, and
long bowl, has a length of 4.25 oral diameters. The oral rim
is ringlike and submerged by the higher rim of the suboral
trough. The suboral trough is very shallow and the diam-
eter of its outer edge is 1.2 oral diameters. The bowl tapers

(12 ) for 0.34 total length, then becomes inverted subconical
(28 ) for 0.29. The transition between lower bowl and
aboral horn is gradual. The horn is conical (15°), is 0.37
total length in length, and has a sharp, pointed free tip.

The wall is thickest suborally (0.12 oral diameter) and
thins evenly below. There are thin laminae with enclosed
minute alveoles in several layers. There are 24 vertical,
continuous, rarely branched and anastomosed ribs with
scattered, irregularly placed fenestrae.

Length, 150 to 225(1.

Rhabdonella valdestriata differs from R. cuspidata in its
longer bowl, lack of aboral swelling, and shortness. It is less
widely conical than R. striata and lacks the distinct suboral
flare of R. spiralis.

Recorded from three stations in the Pacific, as follows:
one (35) in the Pacific equatorial region, one (49) in the
region of South Pacific island fields, and one (63) in the
South Pacific middle latitudes.

There are 3 pump samples and 1 net sample, of which 2
were taken at the surface and 2 at 50 meters. Frequency,
4 per cent in the net sample; 4 to 60 (35.0) loricae in pump
samples.

Temperature: pump samples i7?oi-23?38 (i9?92), net
sample i6?30. Salinity: pump samples 34.59-36.17 (35.13),
net sample 34.88. Density: pump samples 24.36-25.22
(24.76), net sample 25.60. pH: pump samples 8.07-8.27
(8.14), net sample 7.92.

RHABDONELLOPSIS Kofoid and Campbell
Rhabdonellopsis Kofoid and Campbell, 1929, p. 221.

Rhabdonellopsis probably arose from Rhabdonella by the
addition of a knob and lance.

Rhabdonellopsis is a genus of the circumtropical region,
although some species range southward of New Zealand and
northward of Hawaii. Loricae are carried by the Gulf
Stream to considerable latitudes.

Six species are described here.

Rhabdonellopsis apophysata (Cleve) Kofoid and Campbell

Rhabdonellopsis apophysata. Kofoid and Campbell, 1929, p.
221, fig. 420.

The relatively elongated lorica, with stout skirt, ribbed
pedicel, and no fenestrae, has a length of 6.4 oral diameters.
The oral rim is high, ringlike, and erect, and forms the inner
boundary of the suboral trough. The trough is flat, and its
outer rim has a diameter of 1.3 oral diameters. The long
bowl flares (54°) within the upper 0.4 oral diameter. Below
the flaring region the bowl is subconical (15 in the upper
0.35 total length and 32 in the lower section). At its lower
end is the narrow (0.21 oral diameter), cylindrical pedicel,
the length of which is 0.41 total length. At its aboral end is
the expanded skirt, the lower edge of which is irregular, and
from its middle arises the conical (io°) pointed lance.

The wall is thickest just below the suboral trough, where
it reaches 0.12 oral diameter. It becomes much thinner
below. There are thin laminae with enclosed radial, single-

TAXONOMY AND DISTRIBUTION

layered, secondary rectangles, and within these are minute-
alveoles in several layers. On the surface are 20 to 28 sub-
vertical ribs, which continue down the pedicel and over the
skirt. The areas between the ribs are free of fenestrae. The
suboral ring and the lance are hyaline, but the pedicel is
brownish.

Length, 297"..

The ribs are less deflected and the lorica is shorter than
in specimens from other sources.

Rhabdonellopsis apophysata has more ribs than R. longi-
caulis, which species not only is longer, but also is fenestrate
and has a heavier knob. Rhabdonellopsis composita lacks
ribs on the knob, and has fewer ribs on the bowl. Rhabdo-
nellopsis intermedia is shorter, is fenestrate, and has fewer
ribs. In the remaining species the knob is spindle-like.

Recorded from nine stations, seven in the Atlantic and
two in the Pacific, as follows: two (19, 20) in the Sargasso
Sea, two (27, 29) in the Atlantic equatorial region, three
(31, 32, 34) in the Caribbean Sea, and two (35-36, 37) in the
Pacific equatorial region.

There are 12 net samples, of which 3 were taken at the
surface, 3 at 50 meters, and 6 at 100 meters. Maximum fre-
quency, 10 per cent at station 32; other records above mini-
mum (2 to 8 per cent) from stations 27, 32, 34, 35-36; aver-
ages, Atlantic 4.3 per cent, Pacific 3.8 per cent.

Temperature: Atlantic, i8?o8-26?04 (24?8i); Pacific,
1 record only, 27? 12. Salinity: Atlantic, 36.03-37.15 (36.57);
Pacific, 31.68. Density: Atlantic, 23.98-26.06 (25.52); Pacific,
20.20. pH: Atlantic, 8.09-8.30 (8.20); Pacific, 8.28.

Rhabdonellopsis composita (Brandt) Kofoid and Campbell

(Figure 87)

Rhabdonellopsis composita, Kofoid and Campbell, 1929, pp.
222-223, fig. 421.

The rather tall lorica, without ribs on the pedicel, and
with few ribs, stout knob, and thick lance, has a length of
6.05 oral diameters. The oral margin is erect, thin, ringlike,
and higher than the outer edge of the suboral trough. The
suboral trough is concave and the diameter of its rounded-
over outer rim is 1.17 oral diameters. The bowl flares (73 )
within the anterior fifth. Below the flare it tapers (8°) for
0.28 total length, and then becomes subconical (26 ) for
nearly 0.23 total length. The long pedicel is a cylinder (0.25
oral diameter in diameter) with a length of almost 0.42 total
length. At its lower end is the thickened, skirtlike knob.
From the middle of the knob arises the thick lance (0.6 oral
diameter in length).

The wall reaches a maximum thickness of 0.14 oral diam-
eter at the flare and thins down to a tenth that much in the
bowl. There are thin laminae which enclose minute alveoles.
There are 12 vertical, unbranched ribs, which die out below
the middle of the bowl, so that the lower half is nonstriate.
A few (2 to 6) minute fenestrae are found scattered along
each intercosta. The lance has vertical fins. The whole
lorica is colorless, save the knob.

Length, 280 to 353U.

Rhabdonellopsis composita has fewer ribs of shorter length

63

than R. apophysata; it is also fenestrated, and the lance is
fluted.

Recorded from sixteen stations in the Atlantic, as follows:
two (15, 16) in the Gulf Stream, six (17, 18, 19, 20, 20-21,
21) in the Sargasso Sea, seven (22, 23, 24, 26, 27, 28, 29) in
the Atlantic equatorial region, and one (34) in the Carib-
bean Sea.

There are 17 pump and 17 net samples, of which 14 were
taken at the surface, 11 at 50 meters, and 9 at 100 meters.
Maximum frequency, 16 per cent at stations 15, 24; other
records above minimum (2 to 5 per cent) from stations 18,
19, 21; average in net samples, 5.1 per cent; pump samples,
140 loricae taken at station 15, 33 at station 27, 12 at station
34, and 13 at station 24.

Temperature: pump samples i7?5o-28?5i (24?62), net
samples i5?55-26?98 (22^65). Salinity: pump samples
35.22-37.15 (35-54). net samples 35.61-37.15 (36.44). Den-
sity: pump samples 22.84-26.26 (24.20), net samples 23.79-
26.34 ( 2 4-9 2 )- pH: pump samples 8.14-8.37 (8.26), net
samples 7.96-8.32 (8.27).

Rhabdonellopsis intermedia Kofoid and Campbell

Rhabdonellopsis intermedia Kofoid and Campbell, 1929, p. 223,
fig. 424; Marshall, 1934, pp. 650-651, fig. 28.

The short lorica, with flaring conical bowl, tapering shatt,
and deflected ribs, has a length of 4.9 oral diameters. The
oral rim is low and ringlike, and forms the inner boundary
of the concave suboral trough. The diameter of the trough
is 1. 14 oral diameters. The bowl flares (63 ) within the
upper 0.1 oral diameter, then tapers (12°) for 0.31 total
length, and finally becomes inverted subconical (38 ) for
0.8 oral diameter. The pedicel also tapers (8°), the transi-
tion between the lowermost section of the bowl and the
pedicel being gradual. The pedicel also expands a bit above
the knob, so that it is distinctly concave in lateral contour;
it has a length of nearly 0.33 total length. At its lower end
is the modestly expanded knob, from the middle of which
arises the lance (0.5 oral diameter in length), the free tip oi
which is sometimes open.

The wall has a thickness of 0.15 oral diameter suborally
and thins down evenly in the lower bowl. There are thin
laminae which enclose radial, rectangular secondary prisms,
which in their turn enclose minute alveoles in several layers.
The ribs number 16 (24); they are deflected (up to io°) to
the left, and sometimes they branch. They continue on the
pedicel, knob, and lance. The intercostae have 7 to 16 small
fenestrae and strongly developed, though small, prisms.

Length, 222 to 2gS\l.

Rhabdonellopsis intermedia is shorter than R. apophysata,
with fenestrae and fewer ribs. The ribs are continued the
whole length, whereas in R. composita they die out on the
bowl. The pedicel is relatively shorter, as is the whole lorica
actually, than in R. longicaulis.

Recorded from six stations in the Pacific, as follows: two
(45, 46) in the Galapagos region, two (83, 84) in the region
of South Pacific island fields, and two (100, 150) in the
North Pacific trade region.

6 4

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

There are 2 pump and 7 net samples, of which 1 was taken
at the surface, 6 at 50 meters, and 1 each at 100 and 150
meters. Maximum frequency, 11 per cent at station 84;
other records above minimum (2 to 7 per cent) from stations
45, 83, 84; averages, 4.4 per cent and 1.5 loricae in net and
pump samples, respectively.

Temperature: pump samples i4?73~27?46 (2i?oo), net
samples 2i?6o-27?89 (24?7o). Salinity: pump samples
34.27-36.4g (35.38), net samples 34.71-36.42 (35.48). Den-
sity: pump samples 23.11-25.48 (24.59), net samples 22.31-
24.48 (23.79). pH: pump samples 8.12-8.24 (8.18), net
samples 8.12-8.22 (8.16).

Rhabdonellopsis longicaulis Kofoid and Campbell
(Figure 88)
Rhabdonellopsis longicaulis Kofoid and Campbell, 1929, p. 223,

fig. 419.
The greatly elongated lorica, with heavy knob, minute,
sparse fenestrae, and very long pedicel, has a length of 8.0
oral diameters. The oral margin is fairly high, ringlike, and
erect, and is the inner boundary of the concave suboral
trough. The outer margin of the trough is serrate, and its
diameter is 1.24 oral diameters. The bowl flares (52 )
within the anterior 0.32 oral diameter, and then tapers (13 )
for 0.42 total length before becoming subconical (33°) for
0.72 oral diameter. From the lower end of this section arises
the tapering pedicel (5 ), the length of which is 0.42 total
length, and the diameter of which is 0.16 oral diameter. At
the aboral end of the pedicel is the thickened, expanded knob
(0.25 oral diameter in width). From the end of the knob
arises the thin lance, the length of which is 0.6 oral diameter.
The wall reaches nearly 0.1 oral diameter in thickness sub-
orally and lessens to a fraction in the lower bowl and pedicel.
There are thin laminae which enclose minute prisms. The
ribs are few (16), rarely branched within the upper fourth,
continued down to the knob, nearly vertical, decurrent sub-
orally, and not equally spaced. There are 2 to 5 minute
fenestrae scattered in the intercostal spaces; these are the
sites of tiny coccoliths. The knob is brownish, but the rest
of the lorica is translucent.
Length, 350 to 468^.

This large, stately species is easy to distinguish from the
others. The length, few fenestrae, long pedicel, heavy knob,
and few ribs are all characters that separate it.

Recorded from nineteen stations in the Pacific, as follows:
five (45, 46, 75, 76, 77) in the Galapagos region, two (63,
65) in the South Pacific middle latitudes, two (82, 84) in the
region of South Pacific island fields, six (103, 107, 108, 109,
140, 151 ) in the North Pacific trade region, one (135) in the
California region, one (141) in the North Pacific middle
latitudes, and two (152, 153) in the Pacific equatorial region.
There are 21 pump and 12 net samples, of which 16 were
taken at the surface, 11 at 50 meters, and 6 at 100 meters.
Maximum frequency, 40 per cent at station 151; other records
above minimum (2 to 19 per cent) from stations 45, 75, 76,
77, 84, 152; average in net samples, 8.4 per cent; in pump
samples, maximum, 6 loricae at station 45; average, 1.7.

Temperature: pump samples i5?o3-28?o8 (23^43), net
samples n?48-27?52 (22^56). Salinity: pump samples
34.02-35.86 (34.87), net samples 34.02-36.42 (35.43). Den-
sity: pump samples 21.78-25.48 (23.20), net samples 22.34-
26.50 (24.34). pH ; pump samples 8.08-8.47 (8.24), net
samples 7.76-8.21 (8. 11).

Rhabdonellopsis minima Kofoid and Campbell
Rhabdonellopsis minima Kofoid and Campbell, 1929, p. 224,
fig- 4^3-

The rather short lorica, with cornucopia-shaped bowl,
long, tapering pedicel, and spindle-shaped knob, has a length
of 5.0 oral diameters. The oral rim is submerged by the
higher outer edge of the suboral trough, and its diameter is
1.09 oral diameters. The bowl contracts as a cone (13 ) in
the anterior 0.27 total length, and then (25 ) in similar form
and length. The long (0.49 total length), tapering pedicel
(15 ) has a thickened spindle-like knob (0.11 oral diameter
in length), below which is a short lance the length of which
is about equal to that of the knob.

The wall reaches a thickness of 0.1 oral diameter suborally
and progressively lessens lower in the bowl and pedicel.
There are thin laminae which enclose minute alveoles. The
ribs arise a little below the suboral trough, are commonly
bifurcate in the upper part, are decidedly left-turning (as
much as 20 ), continue to the knob and lance, and number
about 16. Fenestrae (6 to 18) are found in the intercostae,
as are also minute prisms.

Length, 177 to 254(^1.

Rhabdonellopsis minima has a conical bowl, more twisted
striae, and spindle-shaped knob, unlike R. intermedia.
Rhabdonellopsis triton is much narrower, with a different
number of ribs and different bowl. The remaining species
have skirted knobs.

Recorded from eight stations in the Pacific, as follows:
one (37) in the Pacific equatorial region, five (52, 54, 61,
62-63, 64) in the South Pacific middle latitudes, and two
(48, 81) in the region of South Pacific island fields.

There are 2 pump and 11 net samples, of which 11 were
taken at the surface and 2 at 100 meters. Maximum fre-
quency, 98 per cent at station 48, minimum at station 52;
average in net samples, 25 per cent; only 3 loricae were in
pump samples.

Temperature: pump samples 26?53~27?i2 (26?82), net
samples io?92-23?63 (i9?53). Salinity: pump samples
31.68-35.82 (33.75), net samples 34.05-36.44 (35.15). Den-
sity: pump samples 20.20-23.50 (21.85), net samples 24.33-
26.06 (24.65). pH: pump samples 8.19-8.28 (8.23), net
samples 8.03-8.27 (8.15).

Rhabdonellopsis triton (Zacharias) Kofoid and Campbell
(Figure 86)

Rhabdonellopsis triton, Kofoid and Campbell. i<)20, p. 224,

fig. 422.
Rhabdonella spiralis, Hofker (part), 1931, pp. 378-381, fig. 67

(for figs. 68-73 see Rhabdonella spiralis).

The relatively short lorica, with conical bowl, swollen,

TAXONOMY AND DISTRIBUTION

65

spindle-shaped knob, long lance, and few ribs and fenestras
has a length of 5.6 oral diameters. The oral rim is a ring-
like cuff which rises above the suboral trough. The suboral
trough is flattened concave, and its outer margin has a diam-
eter of 1.35 oral diameters. The bowl flares (58 ) within
the suboral 0.28, and then becomes inverted, slightly convex
conical (25 ); its length is 0.52 total length. The pedicel is
tubular, with a diameter of only 0.18 oral diameter, anil at
0.S4 total length from the oral margin gives rise to a swollen,
spindle-like knob (0.43 oral diameter in length). From the
lower end arises the lance, the length oi which is 0.4 oral
diameter.

The wall reaches a thickness of almost 0.15 oral diameter
across the suboral trough and gradually thins to a tenth as
much in the pedicel. There are thin laminae and enclosed
minute alveoles in several layers. The ribs number 12; they
are subequidistant and unbranched, and die away before
they reach the knob. There are 2 to 4 minute fenestrae in
each intercostal area. The knob region is brownish, and on
its surface are a number of right-turning striae.
There are 2 macronuclei.
Length, 312U..

The Carnegie loricae have unbranched ribs, few fenestrae,
nonstriate pedicels, and higher oral rims than is usual for the
species from other sources.

Hofker's photograph (fig. 67) shows a spindle-like knob
unlike that of any others of his loricae. This specimen, and
others, are assigned by us to tnton.

Rhabdonellopsis triton has a spindle-shaped knob unlike
the other species, save R. composita. In composita the bowl
is wider, and the ribs are more numerous, are deflected, and
have many fenestrae.

Recorded from three stations, one in the Atlantic and two
in the Pacific, as follows: one (21) in the Sargasso Sea, one
(45) in the Galapagos region, and one (131) in the Cali-
fornia region.

There are 4 net samples, of which 2 were taken at the
surface, 1 at 50 meters, and 1 at 100 meters. Frequency, 3
per cent at station 45; average in the Pacific, 3 per cent.

Temperature: Atlantic, 26? 57; Pacific, 12? 1 2-22?43( 1 8?97).
Salinity: Atlantic, 36.28; Pacific, 33.36-35.26 (34.61). Den-
sity: Atlantic, 23.84; Pacific, 24.30-25. 31 (24.64). pH: At-
lantic, 8.32; Pacific, 8.12-8.32 (8.19).

EPIPLOCYLIDAE Kofoid and Campbell
Epiplocylidae Kofoid and Campbell, 1939, p. 125.
Three genera are included in the family: Epiplocylis, Epio-

rella, and Epicancella. All three are mainly tropical and

occur in the material of this expedition.

EPIPLOCYLIS Jorgensen emended

Epiplocylis, Kofoid and Campbell (part), 1929, pp. 172-173.

Epiplocylis is one of the more advanced genera. The ver-
tical lines suggest derivation from some stem near the Rhab-
donellidae. The cuplike shape remotely resembles that found
in the Petalotrichidae.

Epiplocylis is common in the warm regions of the ocean.
Some species, such as E. undella, are among the most fre-
quent in those areas. Others extend north and south of the
equator into temperate regions, but never reach really cool
waters. Most of the species are circumtropical, with very
little limitation to any of the oceans. Some species occur in
such abundance as to constitute swarms, as do also some
species of Rhabdonella. These swarms in the sea are not
infrequent in some tintinnids, although, as a whole, the
Tintinnoina are more usually found in small percentages.
Areas of abundance usually extend over a wide number of
stations and represent relatively large areas in the ocean.

Seventeen species are described here, one of which is new.

Epiplocylis atlantica Koioid and Campbell
(Figure 72)
Epiplocylis atlantica Kofoid and Campbell, 1929, p. 176, fig.

340.
The short, heavy-set lorica, with wide conical aboral end,
short horn, and low reticulated area, has a length of 1.62 oral
diameters. The oral margin is thinly rounded. The bowl
expands from the rim for 0.55 total length, and has a diam-
eter at the lower level of 1.07 oral diameters. The lower
bowl is inverted conical (93°), and joins the upper bowl
with a rounded angle, the lateral contour of the whole being
distinctly convex. The short (0.22 oral diameter) aboral
horn is conical (28 ) and its free tip is blunted.

The wall has a thickness of 0.067 orru diameter in the sub-
oral thickened region which extends 0.31 oral diameter
below the rim, and is elsewhere about one-third as thick.
There are relatively thick inner and outer laminae which
enclose rather small alveoles. The reticulated aboral region
occupies 0.32 oral diameter, and comparatively few branched
and anastomosing free lines extend above it for 0.15 oral
diameter. The reticulations are deep pits of more or less
subcircular form.
Length, 88|i.

The Carnegie loricae are shorter than those from other
sources.

Epiplocylis atlantica is shorter, wider, and more conical
than E. blanda. In some characters it resembles E. undella,
but it has lateral convexity rather than cylindrical upper
bowl, a full aboral cone, and much finer reticulations.

Recorded from five stations in the Atlantic, as follows:
one (19) in the Sargasso Sea, and four (23, 25, 28, 29) in the
Atlantic equatorial region.

There are 5 net samples, of which 2 were taken at 50

meters and 3 at 100 meters. Frequency, 3 per cent at stations

19, 23; 2 per cent at stations 28, 29; average, 2.5 per cent.

Temperature, i4?6o-27?n (22?73); salinity, 3570-37.05

(36.26); density, 23.62-26.62 (24.92); pH, 7.93-8.29 (8.18).

Epiplocylis blanda Jorgensen emended Kofoid and Campbell
(Figure 73)
Epiplocylis blanda, Kofoid and Campbell, 1929, p. 176, fig. 341;
Marshall, 1934, p. 644, fig. 19.

The tall, cup-shaped lorica, with short free lines and long,

66

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Epiplocylis calyx (Brandt) Jorgensen
Epiplocylis calyx, Kofoid and Campbell, 1929, p. 177, fig. 328.
The short, conical lorica, with thickened suboral wall,
heavy surface reticulation, and sharp aboral horn, has a
length of 1.24 oral diameters. The oral margin is thin and
sharp. The bowl generally has a conical contour (25° in-
creasing to 57 and then to 93°), the angle being least in the
anterior 0.54 total length and gradually greater in the lower
bowl, and quite without sudden changes. The suboral
thickened region (0.28 oral diameter in width) bulges a trifle
beyond the general contour. The aboral horn tapers (20°),
has a length of 0.3 oral diameter, and is sharply pointed at
its free tip.

The wall thickness is 0.06 oral diameter in the wide sub-
oral region and less than a third that much in the remaining
parts of the bowl. The upper 0.24 oral diameter is entirely-
free of surface lines or reticulations. There are only a few
very short free lines, the greater part of the surface being
heavily reticulated with large subcircular, moderately pitted
areas.

Length, 75)1.

Epiplocylis calyx resembles E. labiosa, but is shorter and
wider, with a thinner swollen suboral region, and generally
more conical bowl. Its lower bowl and horn are less abruptly
differentiated than in E. lata, and it has a shorter, wider
bowl and narrower free region than E. exigua.

Recorded from six stations in the Atlantic, as follows: two
(18, 21) in the Sargasso Sea, and four (23, 24, 25, 30) in
the Atlantic equatorial region.

There are 8 net samples, of which 4 each were taken at 50
and 100 meters. Frequency, 5 per cent at station 18; 2 per
cent at stations 24, 30; average, 2 per cent.

Temperature, i4?6o-27?88 (20?84); salinity, 35.61-37.70
(36.41); density, 23.26-26.62 (25.37); P H > 7-93-8-30 (8.02).

Epiplocylis carnegiei, new species
(Plate 1, figure 1 1)
The stout, squat, squarish acorn-shaped lorica has a
length of 2.5 oral diameters. The oral margin is rounded
over, thin, erect, and even. The bowl below is subdivided
into three sections. Of these, the first is a basal segment of
an inverted cone (53°), with a length of 0.54 oral diameter.
Its basal diameter is the widest level of the bowl, 1.53 oral
diameters. Its sides are regular with a bare tendency toward
lateral concavity. The second section is a basal segment of
an inverted truncated cone (25 ) with a length of 0.72 oral
diameter. Its aboral diameter is 1.11 oral diameters. Its
sides are somewhat flattened and regular. Between the above
two conical sections is a broad band with a width of 0.42
oral diameter. Its upper boundary is about 0.26 oral diam-
eter below the oral margin. Its lower boundary contracts

tapering suboral section, has a length of 2.03 oral diameters.
The oral margin is thinly rounded, and there is some suboral
flare (30 ). The long bowl below the flare tapers (2°) for
0.5 total length and then becomes inverted conical (50°);
this section is almost 0.38 total length in length. Its trun-
cated lower end has a diameter of about 0.15 oral diameter.
The aboral horn (0.12 total length in length) is concave
conical (29°), and the free tip is blunt.

The wall is thickest within the flaring section, where it is
0.07 oral diameter; in other parts it is half as much. There
are thin laminae which enclose hyaline material. The reticu-
lated region occupies the aboral 0.56 total length, the upper
third of which has minute circles and short, curved free
lines, the lowermost part having large, more or less circular,
sunken pitlike areas.

Length, 120 to 158^1.

The loricae of this expedition are longer and have fewer
free lines and often shorter aboral horns than usual, and
are much more trim than the one figured by Marshall

(I934)-

Epiplocylis blanda has free lines and a less blunt horn than

E. obtusa. Epiplocylis impensa has different proportions, a
wider aboral cone, and relatively longer horn. Epiplocylis
mucronata is taller and of different proportions, and has a
much longer horn. Epiplocylis acuminata has a very long
horn, and a wide suboral band lacking in blanda. Epi-
plocylis sargassensis has a conical bowl, and the transition
between horn and bowl is more gradual. Once seen, blanda
is scarcely to be confused with the other species.

Recorded from thirty-eight stations, seventeen in the
Atlantic and twenty-one in the Pacific, as follows: one (16)
in the Gulf Stream, five (17, 18, 19, 20-21, 21) in the Sar-
gasso Sea, seven (22, 23, 25, 27, 28, 29, 30) in the Atlantic
equatorial region, four (31, 32, 33, 34) in the Caribbean Sea,
five (47, 68, 71, 78, 80) in the Galapagos region, two (66,
67) in the South Pacific middle latitudes, three (81, 82, 83)
in the region of South Pacific island fields, three (100, 150,
151) in the North Pacific trade region, four (131, 136, 146,

149) in the California region, one (145) in the North Pacific
middle latitudes, and three (152, 153, 154) in the Pacific
equatorial region.

There are 3 pump and 51 net samples, of which 8 were
taken at the surface, 21 at 50 meters, and 25 at 100 meters.

Maximum frequency, 99 per cent at station 131; other
records above minimum (2 to 52 per cent) from stations

16, 17, 19, 20-21, 21, 22, 23, 27, 28, 29, 30, 31, 32, 33, 34, 71,

78, 80, 81, 82, 100, 136, 145, 146, 150, 151, 152, 153, 154;

averages, 9.2 and 13.0 per cent in Atlantic and Pacific net

samples.

Temperature: Atlantic, net samples i4?6o-27?88 (23?29);

Pacific, net samples n?48-27?67 (2^07), pump samples

i3?98-23?88 (i8?83). Salinity: Atlantic, net samples 35.70-

37.05 (36.19); Pacific, net samples 33.24-36.33 (34.61), pump more quickly and assumes the contour of the bowl below.

samples 33.24-35.96 (34.55). Density: Atlantic, net samples This band produces a swollen appearance, and its upper edge

23.84-26.62 (24.80); Pacific, net samples 22.31-26.50 (24.30), separates the bowl into two regions differentiated by wall

pump samples 24.42-25.03 (24.71). pH: Atlantic, net sam- characters. The lowermost of the three sections of the howl

pies 7.93-8.32 (8.23); Pacific, net samples 7.76-8.38 (8.20), is a wide, inverted cone (8o°); its length is 0.42 oral diam-

pump samples 8.23-8.39 (8.31). cter, and is the same as that of the band nearer the oral end

TAXONOMY AND DISTRIBUTION

67

of the howl. This section has full, convex sides formed by
low arcs. It forms an angular junction with the lowest
level of the conical subdivision of the bowl above it. The
aboral horn is a short, conical peg (30 ), the length of which
is only 0.16 oral diameter.

The wall is thickened in the suboral region and the band.
Its greatest thickness in these places is 0.14 oral diameter. In
the bowl it is much thinner, being less than 0.04, more or
less subuniformly at all levels. The wall is filled with small,
irregularly arranged secondary prisms which crowd upon
one another so that no two are shaped alike. In the thinner
region of the bowl there are fewer layers of prisms than in
the superstructure. The outer wall is differentiated into two
zones by the patterning. Of these, the anterior one is the
narrower band. This begins at the upper edge of the suboral
band and continues to the oral margin. It is made up of
faint, roughly hexagonal prisms, about 10 in number in a
vertical line and approximately 60 in number across the face
of the bowl. They increase in size and prominence as they
reach the suboral ledge, and an exact plane of division be-
tween them and the coarse reticulum below becomes difficult.
The coarse surface reticulum which incrusts the posterior
0.85 of the bowl is made up of a prevailingly pentagonal to
hexagonal meshwork. The nearly uniform framework which
surrounds the depressed areas is heavy and thick, and gives
the lorica a clumsy appearance. There are 12 to 16 meshes
in a vertical line and about 20 to 26 across the bowl. The
mesh continues over the basal part of the aboral horn, but
the free end is devoid of extra structure. The lumen of the
upper bowl is a cylinder with a length of about 0.26 oral
diameter. Below this level it is a basal segment of a cone
(about 6o°) with a length of nearly 0.26 oral diameter. It
then follows the outer contour reasonably closely to the
lower end of the bowl. The horn is solid.

Length, bowl 10011, horn 12. 5|A; diameter, oral 56.211,
maximum 8ij.i, lower bowl 63U.; wall thickness, 8(x.

The shape of the lower bowl varies. Though having a
general tendency to be squarish, it may narrow and be more
or less conical. Such loricae recall Epiplocylis semireticulata.
The thickened zone is wider in some loricae than in others,
and the inner side of the throat, though typically vertical,
may flare outward. The surface reticulations are thicker-
walled and coarser in some individuals than in the general
run.

Epiplocylis carnegiei is close to E. semireticulata, but dif-
fers from that species in more contracted oral aperture, rela-
tively shorter aboral horn, and less coarse reticulum. Loricae
from this collection have greater suboral contraction than the
loricae figured by Brandt (1906, pi. 58, fig. 9) and their
lower ends are squarish in form; otherwise they are similar.
This species differs from E. acuminata in its more definitely
oblong shape and short horn as well as in its finer and
heavier reticulum. The aboral horn of acuminata is nearly
0.33 total length in length.

Epiplocylis semireticulata was described by Biedermann
(1893) and has had a checkered career, which has been
summed up by Kofoid and Campbell (1929). In the present
report it is considered as distinct from E. acuminata, with

which the latest authors have put it. Perhaps Brandt'-,
lorica belongs to carnegiei and not to semireticulata; the
djfferences may be only minor ones within the limits of a
single species.

Recorded from five stations in the Atlantic, as follows:
two (3, 4) in the Atlantic drill, and three (14, 15, 16) in
the Gulf Stream.

There are 4 pump and 5 net samples, oi which 2 were
taken at the surface, 3 at 50 meters, and 4 at 100 meters.
Maximum frequency, 7 per cent at station 3; other records
above minimum ( 2 to 3 per cent) trom station 16; average
in net samples, 3.6 per cent.

Temperature: pump samples 14^02-24^81 (20?52), net
samples i3?37~23?64 (i6?35). Salinity: pump samples
35.59-36.48 (36.21), net samples 35.88-36.41 (36.02). Den-
sity: pump samples 24.47-26.66 (25.49), net samples 24.84-
27.01 (26.40). pH: pump samples 8.06-8.23 (8-i6), net
samples 8.10-8.23 (8.15).

Type locality, station 3, at 50 meters; latitude 44 00'
north, longitude 36 10' west.

Epiplocylis constricta Kofoid and Campbell

Epiplocylis constricta Kofoid and Campbell, 1929, p. 177, fig.
333; Marshall, 1934, p. 643.

The fairly short, rotund lorica, with short free lines and
narrow aboral horn, has a length of 1.72 oral diameters.
The oral margin is thin and rounded. The bowl is dis-
tinctly rotund, expanding from the rim (12 ) for 0.64 total
length and reaching 1.09 oral diameters at that level. The
aboral part contracts convexly (33 increasing to 8o° pos-
teriorly) and the aboral horn arises from its lowermost end.
The horn is conical (18 ) and nearly 0.55 oral diameter in
length, and has a sharp free tip.

The wall has a maximum thickness of 0.07 oral diameter
in the suboral region and thins to less than a third as much
aborally. There are thin, homogeneous laminae with en-
closed tiny alveoles. The reticulated region occupies the
aboral half of the bowl, and above that level the free lines
arise. The short free lines are deflected (15 to 50 ) toward
the left; the reticulations are thick-walled, subcircular, and
often overlapping, and on the upper horn are heavy vertical
lines.

Length, 93 to ii2p.

Epiplocylis constricta has shorter tree lines than E. de-
flexa, and no oral flare. In E. inconspicuata the deflected
free lines extend much farther on the wider bowl. Other
related species have subvertical free lines.

Recorded from twenty-six stations in the Pacific, as fol-
lows: one (54) in the South Pacific middle latitudes, eight
(82, 83, 84, 85, 157, 158, 159, 160) in the region of South
Pacific island fields, four (100, 109, 140, 150) in the North
Pacific trade region, three (113, 142, 145) in the North
Pacific middle latitudes, five (137, 146. 147, 148, 149) in the
California region, and five (152, 153, 154, 155, 156) in the
Pacific equatorial region.

There are 15 pump and 25 net samples, of which 8 were
taken at the surface, 16 at 50 meters, and 16 at ioo meters.

68

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Maximum frequency, 64 per cent at station 155; other records
above minimum (2 to 44 per cent) from all stations except
54; average in net samples, 15.3 per cent; in pump samples
there were 1 to 6 loricae, average 1.7.

Temperature: pump samples i4?32-27?73 (24?i5), net
samples i8?74~28?6o (25?i6). Salinity: pump samples
34.50-35.04 (34.77), net samples 34.18-36.49 (34.93). Den-
sity: pump samples 22.29-25.7s (23.45), net samples 22.65-
25.37 ( 23.54). pH: pump samples 7.87-8.47 (8.34), net
samples 7.93-8.44 (8.28).

Epiplocylis deflexa Kofoid and Campbell
Epiplocylis deflexa Kofoid and Campbell, 1929, p. 178, fig. 334;
Marshall, 1934, p. 645, fig. 20.

The moderately large lorica, with convex bowl, conical
aboral horn, and strongly deflected free lines, has a length of
1.67 oral diameters. The oral margin is thick, but sharp
and erect. The wide bowl increases regularly from the rim
to 1. 1 7 oral diameters at 0.5 oral diameter below the rim.
The lower bowl is convex conical (20 increasing to 74 °
and then to 93 ), being least in its anterior half and grad-
ually wider below. The aboral horn is narrow conical (23 )
with a length of 0.28 oral diameter, and is sharply pointed
at its free tip.

The wall is angular in section. Its thickness (maximum
0.14 oral diameter) reduces the cavity to 0.89 oral diameter
near 0.17 below the rim, and is gradually reduced by thin-
ning lower down. There are relatively thick laminae with
large, rectangular secondary areas; these last, in turn, enclose
minute primary alveoles. The reticulated aboral region
occupies the lowermost 0.67 oral diameter. The reticula-
tions are large, subcircular, deeply pitted areas. The free
lines are strongly deflected to the left (up to 6o°) and then
extend upward to within the anterior 0.25 of the bowl. On
the upper end of the horn there are strong vertical ridges.
Length, 88 to 11 $\i.

The Carnegie material agrees closely with Marshall's.
Epiplocylis deflexa resembles E. constricta, but the free
lines are more strongly deflected and longer, the wall is
thicker, with secondary areas, and the horn is not so long.
Epiplocylis pacifica is longer and more trim, with shorter
free lines which are vertical rather than deflected. Epiplocylis
inconspicuata has a wider (not flattened) bowl with sub-
cylindrical upper section.

Recorded from eleven stations in the Pacific, as follows:
four (45, 46, 47, 80) in the Galapagos region, four (48, 82,
85, 95) in the region of South Pacific island fields, two (99,
153) in the Pacific equatorial region, and one (113) in the
North Pacific middle latitudes.

There are 2 pump and 12 net samples, of which 3 were
taken at the surface, 5 at 50 meters, and 6 at 100 meters.
Maximum frequency, 21 per cent at station 47; other records
above minimum ( 2 to 18 per cent) from stations 45, 46, 48,
82, 95, 99, 113, 153; average in net samples, 6.6 per cent.

Temperature: pump samples 23: 58-28^05 (25^81 ), net
samples 2i?69-28?74 (24^45). Salinity: pump samples
34.40-36.21 (35.30), net samples 34.66-36.44 (35.49). Den-

sity: pump samples 21.95-24.70 (23.32), net samples 22.43-
24.48 (23.86). pH: pump samples 8.18-8.39 ( 8 - 28 )> net

samples 8.12-8.28 (8.19).

Epiplocylis exigua Kofoid and Campbell

Epiplocylis exigua Kofoid and Campbell, 1929, p. 178, fig. 337;

Marshall, 1934, p. 643, fig. 17.
The rather small lorica, with subcylindrical upper and
conical lower bowl, thickened suboral wall, well reticulated
surface, and gradually differentiated aboral horn, has a
length of 1.83 oral diameters. The oral margin is thick.
The bowl is convex subcylindrical in the anterior 0.65 oral
diameter, and conical below (53 in the upper two-thirds
and 8o° in the lower part). The aboral horn is gradually
formed, being conical (16 ), with a length of 0.38 oral
diameter, and blunt at its free tip.

The wall has a thickness of 0.1 oral diameter in the
thickened zone, which occupies the anterior 0.41 oral diam-
eter; its thickness is reduced to a third as much in the
remaining part of the bowl. There are rather thick laminae
which enclose hyaline material. The reticulated region
occupies about 0.75 of the length of the bowl; the reticula-
tions are large, squarish, pitted areas. A few vertical free
lines occur, and on the upper end of the horn are upright
lines.

Length, 74 to 88(.i.

This species intergrades in some characters with Epi-
plocylis labiosa, as Marshall (1934) points out, but it may
be distinguished by the fuller lower bowl, the more gradually
differentiated horn, and the higher level of the reticulated
area. Its horn is not so long as that of E. lata, and the bowl
and horn are less distinct; the reticulated region is typically
higher in lata.

Recorded from four stations in the Pacific, as follows: one
(78) in the Galapagos region, one (83) in the region of
South Pacific island fields, one (113) in the North Pacific-
middle latitudes, and one (136) in the California region.

There are 3 pump samples and 1 net sample, of which 1
was taken at 50 meters and 3 at 100 meters. Frequency, 12
per cent at station 113; other records minimum.

Temperature: pump samples i8?87~27?46 (22?46), net
sample 2i?74- Salinity: pump samples 35.02-36.49 (35.56),
net sample 34.66. Density: pump samples 23. 71-25. n
(24.63), net sample 24.06. pH: pump samples 8.14-8.39
(8.25), net sample 8.23.

Epiplocylis exquisita Kofoid and Campbell
Epiplocylis exquisita Kofoid and Campbell, 1929, p. 179, fig.

342-
The moderately tall, wide lorica, with convex subcorneal
bowl, has a length of 1.25 oral diameters. The oral margin
is thin and fairly sharp. The bowl is very wide, and convex
subconical (18 increasing to 48 and then to 90 ), the
angle being least in the anterior 0.46 oral diameter and
greatest in the aboral 0.3 oral diameter. The aboral horn is
narrow conical (18 ), and 0.33 oral diameter in length, with
a pointed free tip.

TAXONOMY AND DISTRIBUTION

69

The wall has a maximum thickness of 0.04 oral diameter
in the anterior part of the bowl, and two-thirds as much in
the lower bowl. There are exceedingly thin laminae with
enclosed hyaline matter. The anterior half of the bowl is
free of markings. The free lines are short, anastomosing,
thin, subvertical lines, and the reticulations are large, thick-
walled areas of subcircular form. Most of the horn is
hyaline.

Length, 93 to 1 1 oil.

Epiplocylis exquisita has a wider, shorter bowl than E.
blanda, thinner walls, shorter free lines, and more sharply
differentiated horn than E. undella, much less rotund bowl
than E. pacified, and shorter bowl than E. impensa.

Recorded from four stations, one in the Atlantic and three
in the Pacific, as follows: one (25) in the Atlantic equatorial
region, two (35, 35-36) in the Pacific equatorial region, and
one (100) in the North Pacific trade region.

There are 4 net samples, of which 1 was taken at the
surface, 2 at 50 meters, and 1 at 100 meters. Frequency, 39
per cent at station 100; 4 to 7 per cent at stations 25, 35-36;
average in the Pacific, 21.5 per cent.

Temperature: Atlantic, i4?6o;Pacific,i6?30-27?67(2i?98).
Salinity: Atlantic, 35.70; Pacific, 34.71-34.88 (34.79). Den-
sity: Atlantic, 26.62; Pacific, 22.31-25.60 (23.95). pH: At-
lantic, 7.93; Pacific, 7.92-8.21 (8.06).

Epiplocylis impensa Kofoid and Campbell

Epiplocylis impensa Kofoid and Campbell, 1929, p. 180, fig. 343.

The fairly tall, tapering, goblet-shaped lorica, with strong
angles, low reticulated region, and thick aboral horn, has a
length of 1.44 oral diameters. The oral margin is sharp-
edged and erect. The bowl is conical (io°) in the upper
0.56 oral diameter, the diameter at the lower end being 0.93
oral diameter. Below this level it contracts (33 in the
upper part increasing to 83 below); the upper part is 0.94
oral diameter in length. The lateral contour of the whole
is distinctly angled at important levels of change. The aboral
horn is conical (23 ), and blunt at the free tip.

The wall reaches a thickness of a little more than 0.06
oral diameter in the suboral region; in the lower bowl it is
about half as much, or less. There are thin laminae with
enclosed alveoles. The reticulated region occupies about 0.5
oral diameter and the free lines extend subvertically for as
much as 0.6 oral diameter. The reticulations are moderately
pitted and subpentagonal to hexagonal, and on the horn
free vertical lines are continued for 0.75 of its length.

Length, 115 to 126"..

Epiplocylis impensa has angled instead of convex lateral
contour, and a longer horn and much coarser reticulations
than E. atlantica, which is otherwise much like it. Epiplocylis
lineata, its neighbor, is conical, with a long horn and very
long free lines. Epiplocylis blanda is narrower, with a longer
free region and finer reticulations.

Recorded from two stations in the Atlantic, as follows:
one (19) in the Sargasso Sea and one (27) in the Atlantic
equatorial region.

There are 3 net samples, 2 of which were taken at 50

meters and 1 at 100 meters. Frequency, 8 per cent al
station 27.

Temperature, [8?o8-26?04 (23?i4); salinity, 36.03-37.15
(36.47); density, 23.98-26.06 (24. hi); pH, 8.09-8.30 (8.22).

Epiplocylis inconspicuata Kofoid and Campbell

Epiplocylis inconspicuata Kofoid ami Campbell, 1929, pp. 180-
181, fig. 326.

The short, wide lorica, with conical bowl and narrow free
region, has a length of 1.2 oral diameters. The oral margin
is sharp-edged. The bowl expands with distinct lateral con-
cavity to a diameter of 1.04 oral diameters near 0.18 oral
diameter below the rim. The swollen band has a width o!
approximately 0.27 oral diameter. The bowl below tins
lower level contracts (44 increasing to t 15 ) , being least
in the anterior 0.38 oral diameter and thence, with sharp
angular change, greater. The aboral horn is narrow conical
(26°) and only 0.2 oral diameter in length, and has a
sharply pointed free tip.

The wall is thickest in the swollen region, where it is over
0.07 oral diameter; lower down in the bowl it is less than a
third as much. There are extraordinarily thin laminae
which enclose pallid, minute alveoles. There are no free
lines, the upper region being 0.15 oral diameter in width.
The reticulations are coarse, subequal, deeply pitted, and
squarish to rounded.

Length, 72(1.

Epiplocylis inconspicuata differs from E. infiata in having
a narrower free region, less suboral thickening, and less
width.

Recorded from two stations (65, 67) in the South Pacific
middle latitudes, in 2 net samples taken at 100 meters.
Frequency, minimum.

Temperature, i5?03-i6?36 (15^69); salinity, 34.30-34.70
(34.50); density, 25.44-25.45 (25.445); pH, 8.09-8.10 (8.095).

Epiplocylis infiata Kofoid and Campbell
Epiplocylis infiata Kofoid and Campbell, 1929, p. 181, fig. 329.

The short, distinctly wide lorica, with short free region,
has a length of 1.0 oral diameter. The oral margin is thin
and sharp. The bowl is subcylindrical in the anterior 0.16
oral diameter, then swollen to a diameter of 1.07 oral diam-
eters within a band 0.22 oral diameter in width, and grad-
ually contracts in the remaining region (50° increasing to
105 ), the angle being least in the upper two-thirds of the
section. The aboral horn is narrow conical (23 ) and only
0.16 oral diameter in length, and has a sharp free tip.

The wall is thickest in the wide band, where it reaches
about 0.09 oral diameter; elsewhere it is reduced to a third as
much, or less. There are extremely thin laminae, between
which are packed minute alveoles. The surface reticulations
are lacking in the subcylindrical section. There are a few
scattered, subvertical, short free lines, and the reticulations
are large subpentagons or hexagons.

Length, 77 to 85U.

7 o

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Epiplocylis inflata is much like E. laac\manni, but its
horn is shorter and the lower bowl is fuller. It is much
wider and relatively shorter, and has a wider free region
than E. calyx. It only remotely resembles E. acuminata,
which has a longer bowl and horn and different pro-
portions.

Recorded from station 23 in the Atlantic equatorial region,
in a net sample taken at 50 meters. Frequency, 2 per cent.

Temperature, 20?99; salinity, 36.04; density, 25.30; pH,
8.14.

Epiplocylis labiosa Kofoid and Campbell

(Figure 74)

Epiplocylis labiosa Kofoid and Campbell, 1929, p. 182, fig. 338.

The subconical, rather elongated lorica, with short reticu-
lated region and thick suboral area, has a length of 1.87 oral
diameters. The oral margin is thin and erect. The acorn-
shaped bowl expands (45°) to 1.17 oral diameters at 0.22
oral diameter below the rim and then contracts below the
thickened zone (0.38 oral diameter in width) as a segment
of inverted flat-sided cone (30 ) with a length of 0.43 oral
diameter; its diameter at the lower end is 0.81 oral diameter.
Below this section the bowl becomes more widely conical
(73 ), and the diameter at the truncate lower end is only
0.16 oral diameter. The aboral horn is 0.54 oral diameter
in length and narrow conical (15°). Its free tip is sharp.

The wall has a thickness of 0.16 oral diameter in the
thickened suboral zone, but elsewhere is reduced to three-
tenths as much. There are extremely thin laminae, which
enclose many layers of very small alveoles. The reticulated
region is deeply pitted and coarse, and covers the lowermost
conical section of the bowl; above it is a faint network of
large, virtually hexagonal areas which reach to just below
the thick suboral region; the vertical lines of the mesh are
stronger than the horizontal ones.

Length, 73 to 85^1.

The Carnegie loricae are shorter and have a finer mesh
than is usual for this species.

Epiplocylis labiosa is longer and narrower, with a shorter
free zone and a thicker suboral band than E. calyx. Epi-
plocylis exigua is shorter and wider and has a lower zone of
reticulations. Epiplocylis lata has a more cylindrical upper
bowl, wider aboral cone, and even coarser reticulum.

Recorded from thirty stations, ten in the Atlantic and
twenty in the Pacific, as follows: four (19, 20, 20-21, 21) in
the Sargasso Sea, six (22, 23, 24, 25, 29, 30) in the Atlantic
equatorial region, four (82, 85, 87, 93) in the region of
South Pacific island fields, ten (100, 101, 102, 103, 104, 105,
106, 107, 150, 151) in the North Pacific trade region, four
( 137, 147, 148, 149) in the California region, and two ( 153,
154) in the Pacific equatorial region.

There are 26 pump and 20 net samples, of which 15 were
taken at the surface, 17 at 50 meters, and 14 at 100 meters.
Maximum frequency, 6 per cent at station 21; other records
above minimum (2 to 3 per cent) from stations 19, 22, 137,
153, 154; averages, 1.6 and 1.7 per cent in Atlantic and
Pacific net samples; none of the pump samples had more
than 3 loricae.

Temperature: Atlantic, pump samples i4?6o-27?56(22?79),
net samples i 4 ?6o-27?88 (23?02); Pacific, i8?28-25?8i
(22?^) and 23?i5~28?74 (25?95), respectively. Salinity:
Atlantic, pump samples 35.70-36.21 (35.80), net samples
35-7°-37- I 5 (36-29); Pacific, 34.39-36.32 (35-16) and 34.42-
35.12 (34.81), respectively. Density: Atlantic, pump samples
23.36-26.62 (24.65), net samples 23.26-26.62 (24.73); Pacific,
21.95-24.26 (23.26) and 22.98-24.77 (24.03), respectively.
pH: Atlantic, pump samples 7.93-8.31 (8.17), net samples
7.93-8.37 (8.21); Pacific, 8.21-8.34 ( 8 - z6 ) and 7-93~ 8 -3 8
(8.23), respectively.

Epiplocylis lata Kofoid and Campbell
(Figure 78)
Epiplocylis lata Kofoid and Campbell, 1929, p. 182, fig. 336.

The short lorica, with greatly thickened suboral band and
abruptly contracted aboral end, has a length of 2.2 oral
diameters. The oral rim is the rounded edge of the thick-
ened suboral region. The bowl expands rapidly (55°) for
0.24 oral diameter to 1.07 oral diameters. Below this level it
tapers (14 ) for 1.0 oral diameter, the diameter at the lower
end being 0.92 oral diameter. The aboral region is convex
conical (8o°) for about 0.5 oral diameter. At its lower end
is the sharply pointed, conical (16°) aboral horn, the length
of which is 0.6 oral diameter.

The wall thickens to 0.16 oral diameter in the suboral
region (the width of the band reaches 0.52 oral diameter),
and is elsewhere less than three-tenths as much. The reticu-
lated area extends to a little below the thickened zone and
is made up of large rectangular to pentagonal areas; some
are even subcircular, and all are deeply pitted. The horn
has long vertical ridges on its upper part.

Length, 82 to nop:.

A few of the Carnegie loricae are longer than is usual for
this species.

Epiplocylis lata is more abruptly rounded aborally and its
aboral horn, which is distinctly spinelike, is set off more
suddenly from the lower bowl than in E. calyx, E. exigua, or
E. labiosa. In the Carnegie collection it is not easily distin-
guished from labiosa.

Recorded from fourteen stations in the Pacific, as follows:
six (45, 46, 47, 75, 78, 80) in the Galapagos region, four
(48, 82, 84, 85) in the region of South Pacific island fields,
and four (54, 61, 64, 65) in the South Pacific middle lati-
tudes.

There are 7 pump and 16 net samples, of which 7 were
taken at the surface, 11 at 50 meters, and 5 at 100 meters.
Maximum frequency, 44 per cent at station 45; other records
above minimum (2 to 5 per cent) from stations 46, 47, 48,
54, 82, 84, 85; average in net samples, 7.9 per cent; in pump
samples there were 1 to 33 loricae, average 10.8.

Temperature: pump samples i4?03-23?30 (20?82), net
samples io?92-27?89 (22?23). Salinity: pump samples
33-9 6 -35-9 6 (3v°9)- net samples 34.05-36.44 (35.55)- Den-
sity: pump samples 24.11-25.40 (24.52), net samples 23.38-
26.06 (25.23). pH: pump samples 8.05-8.16 (8.13), net
samples 8.03-8.23 (8.15).

TAXONOMY AND DISTRIBUTION

71

Epiplocylis mucronata (Zacharias) Kofoid and Campbell

(Figures 75, 76)

Epiplocylis mucronata, Kofoid and Campbell, 1929, p. 183, fig.
346.

The tall, conical lorica, with gradually differentiated aboral
born and wide free region, has a length of 2.00 to 2.92 oral
diameters. The oral margin is sharp-edged. The long bowl
tapers as much as 10° for 1.2 oral diameters and shows some
lateral convexity. The lower bowl contracts (45° to 65 )
for r.13 oral diameters, the change in diameter between
upper and lower bowls being gradual. The aboral horn is
gradually differentiated out of the lower bowl, is conical
(22 to 30 ), and has a length of 0.16 to 0.30 oral diameter;
its free tip is sharp.

The wall reaches a maximum thickness of 0.1 oral diam-
eter suborally; elsewhere it is reduced to less than one-fifth
as much. The laminae, with enclosed alveoles, are thin.
The upper bowl is entirely free of lines; subvertical, thick,
and unbranched free lines creep to its lower edge from the
reticulated region below. The reticulations are coarse, thick-
walled, heavy, and not deeply pitted, and on the horn are
extended as heavy vertical ridges.

Length, 175 to 18511.

The Carnegie loricae are much more gradually contracted
at the transition of lower bowl and horn than is typical;
there are also strong free lines.

Epiplocylis mucronata differs from E. obtusa, in which
there is a saccular bowl and short, wide horn. It differs
from E. blanda in larger size, more taper, and heavier reticu-
lum. Epiplocylis sargassensis is generally more conical and
wider in the aboral region. Epiplocylis acuminata has a
wider suboral thickening and spikelike horn, as well as
different mesh.

Recorded from six stations, three each in the Atlantic and
the Pacific, as follows: two (28, 29) in the Atlantic equa-
torial region, one (31) in the Caribbean Sea, one (47) in the
Galapagos region, and two (64, 65) in the South Pacific
middle latitudes.

There are 8 net samples, of which 2 were taken at the
surface, 1 at 50 meters, 4 at 100 meters, and 1 at 1000
meters. Maximum frequency, 54 per cent at station 47;
other records above minimum (2 to 3 per cent) from stations
28, 31, 65; average in Pacific, 14.7 per cent.

Temperature: Atlantic, 22?56-26?79 (23?94); Pacific,
3?98-23?88 (i5?87). Salinity: Atlantic, 36.31-36.63 (36.51);
Pacific, 34.30-35.96 (34.79). Density: Atlantic, 23.79-25.22
(24.80); Pacific, 24.33-27.23 (25.36). pH: Atlantic, 8.19-
8.26 (8.22); Pacific, 7.76-8.23 (8.06).

Epiplocylis pacifica Kofoid and Campbell

Epiplocylis pacifica Kofoid and Campbell, 1929, p. 184, fig. 335.

The fairly tall lorica, with moderately convex conical bowl,
short free lines, and long aboral horn, has a length of 1.83
oral diameters. The oral margin is thin and erect. The
bowl expands from the margin to 1.08 oral diameters at 0.3
oral diameter below the rim, and gradually contracts (20

then 80 c ' in the aboral 0.5 oral diameter) to the aboral end.
The aboral horn is narrow conical (12°), nearly 0.46 oral
diameter in length, and sharply pointed at its free tip.

The wall reaches a thickness of 0.09 oral diameter across
the thickest part, which is within a band 0.67 oral diameter
in width; the lower parts oi the bowl are less than a third
as much. The upper region ol the bowl, corresponding to
the thickened zone in width, is free ol reticulations. The
reticulations of the lower region arc large, commonly sub-
pentagonal, with thick walls, and modestly pitted, and have
short, unbranched, deflected (20 ) free lines.

Length, 105 to 1 15(1.

Epiplocylis pacifica resembles E. constricta, but its bowl is
decidedly less convex, its horn is longer, and the free lines
are much shorter and less oblique. In general form it is
close to E. lata, but it is obviously dillerent in the more
diffuse suboral thickening, and in the character of the reticu-
lum and its distribution. In some ways it is like E. undella,
but it is less wide, with more sharply differentiated horn,
and with shorter, deflected free lines.

Recorded from sixteen stations in the Pacific, as follows:
three (35, 36, 99) in the Pacific equatorial region, one (62)
in the South Pacific middle latitudes, two (77, 78) in the
Galapagos region, four (81, 82, 83, 159) in the region of
South Pacific island fields, three (101, 102, 103) in the North
Pacific trade region, and three (133, 134, 137) in the Cali-
fornia region.

There are 15 pump and 7 net samples, of which 11 were
taken at the surface, 5 at 50 meters, and 6 at 100 meters.
Maximum frequency, 7 per cent at station 77; other records
above minimum (2 to 5 per cent) from stations 78, 159;
average in net samples, 3.1 per cent; in pump samples there
were 1 to 4 loricae.

Temperature: pump samples i3?28-27?93 (24?47), net
samples i4?33-28?6o (24^80). Salinity: pump samples
31.62-36.49 (34.98), net samples 34.91-36.03 (35.70). Den-
sity: pump samples 20.34-24.98 (22.77), net samples 22.77-
26.06 (23.79). pH: pump samples 8.06-8.39 (8.25), net
samples 7.88-8.37 (8.18).

Epiplocylis sargassensis (Brandt) Kofoid and Campbell

Epiplocylis sargassensis, Kofoid and Campbell, 1929, p. 185,
fig- 331-

The moderately elongated, generally conical lorica, with
full aboral region and broad horn and lacking free lines, has
a length of 1.6 oral diameters. The oral margin is thin and
rounded. The bowl tapers (16 ) in its upper 0.52, the
diameter at the lower end of this upper section being 0.5
oral diameter. The lower bowl is subcorneal (55°), and
from its lower end the aboral horn is gradually differen-
tiated; this latter becomes narrow conical (20 ), its length
is 0.26 oral diameter, and its free tip is blunted.

The wall reaches as much as 0.03 oral diameter at its
thickest, which is near the oral rim. In the middle and
lower bowl it is about two-thirds as much. There are thin
laminae and enclosed minute alveoles. There are no free
lines, but the reticulated region occupies the lower 0.83 oral

72

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

diameter. The reticulations are subcircles, large and heavy-
walled, and smaller circles occur with the large ones. On
the upper horn are heavy vertical lines. All the reticulations
are deeply pitted.

Length, 130^1.

Epiplocylis sargassensis has a fuller lower bowl and
shorter horn than E. mucronata; it also has no free lines. It
differs from E. blanda in similar respects. It has a less
baggy bowl and longer horn than the peculiar species E.
obtusa.

Recorded from twenty stations, nine in the Atlantic and
eleven in the Pacific, as follows: three (19, 20, 21) in the
Sargasso Sea, four (23, 28, 29, 30) in the Atlantic equatorial
region, two (31, 34) in the Caribbean Sea, seven (134, 135,
136, 146, 147, 148, 149) in the California region, two (144,
145) in the North Pacific middle latitudes, and two (150,
151) in the North Pacific trade region.

There are 5 pump and 18 net samples, of which 2 were
taken at the surface, 4 at 50 meters, and 17 at 100 meters.
Maximum frequency, 13 per cent at station 29; other records
above minimum (2 to 12 per cent) from stations 20, 21, 28,
30, 31, 34, 136, 146, 147, 149, 151; averages in net samples
6.8 and 4 per cent in the Atlantic and Pacific, respectively.

Temperature: Atlantic, net samples 20?99~27?88 (24?i9);
Pacific, net samples i8?87-25?95 (20^56), pump samples
i6?58— ig?i6 (i8?o6). Salinity: Atlantic, net samples 36.04-
37.05 (36.45); Pacific, net samples 34.02-35.04 (34.70), pump
samples 34.32-34.88 (34.62). Density: Atlantic, net samples
23.26-25.76 (24.67); Pacific, net samples 22.34-25.09 (24.38),
pump samples 24.48-25.11 (24.90). pH: Atlantic, net
samples 8.14-8.32 (8.22); Pacific, net samples 8.26-8.39
(8.32), pump samples 8.31-8.37 (8.34).

Epiplocylis undella (Ostenfeld and Schmidt) Jorgensen
emended Kofoid and Campbell

(Figures 71, 77)

Epiplocylis undella, Kofoid and Campbell, 1929, p. 185, fig. 345;
Marshall, 1934, p. 645, fig. 18.

The rather large, wide lorica, with conical aboral end,
tapering horn, short reticulated zone, and short free lines,
has a length of 2.06 to 2.28 oral diameters. The oral margin
is thin, rounded, and angular. The bowl is usually cylin-
drical in the anterior 1.0 oral diameter, with a diameter of
as much as 1.23 oral diameter at the lower end in unusually
swollen individuals. The lower bowl is distinctly convex
conical (72 to 8o°). The aboral horn tapers (16 to 18 ),
is sharply pointed, and has a length of 0.50 to 0.53 oral
diameter.

The wall is relatively thick, reaching 0.11 oral diameter
through the upper anterior bowl, and gradually becoming
less than half as much lower down. There are thick laminae
which enclose small, circular alveoles in two to five layers.
The upper 1.0 oral diameter is free of lines and reticulations.
The reticulated zone occupies approximately the lower two-
thirds of the lower bowl, and the free lines, on occasion,
reach the lower end of the upper bowl. The reticulations
are subcircular, rather large, in 4 or more rows, and deeply

pitted. The free lines are subvertical, relatively few, and
mostly unbranched.

Length, 103 to i8o[.i.

The Carnegie loricae are longer (168 to i8op) than usual
(as low as 103(1), with convex sides and swelling, and with
thick laminae.

Epiplocylis undella resembles E. blanda in general shape,
but may be distinguished by the much shorter upper region
in which there are no free lines; it is generally wider. Epi-
plocylis mucronata has a very much longer bowl and coarser
reticulations.

Recorded from seventy-six stations, nineteen in the At-
lantic and fifty-seven in the Pacific, as follows: two (15. 16)
in the Gulf Stream, six (17, 18, 19, 20, 20-21, 21) in the Sar-
gasso Sea, nine (22, 23, 24, 25, 26, 27, 28, 29, 30) in the
Atlantic equatorial region, two (33, 34) in the Caribbean
Sea, seven (37, 38, 99, 152, 153, 155, 156) in the Pacific equa-
torial region, nine (45, 46, 47, 69, 70, 71, 77, 78, 80) in the
Galapagos region, five (62, 62-63, 65, 66, 67) in the South
Pacific middle latitudes, seventeen (81, 82, 84, 85, 86, 89, 90,
91, 92, 93, 95, 96, 97, 157, 158, 159, 160) in the region of
South Pacific island fields, nine ( 100, 101, 105, 107, 109, 139,
140, 150, 151) in the North Pacific trade region, eight (131.
133, 134, 135, 136, 137, 146, 148) in the California region,
and two (142. 14s) in the North Pacific middle latitudes.
Epiplocylis undella and Eutintinnus lusus-undae are perhaps
the most abundant species of Tintinnoina in the warmer
regions of the ocean; either may be expected in almost any
plankton catch in tropical waters.

There are 84 pump and 64 net samples, of which 42 were
taken at the surface, 48 at 50 meters, and 58 at 100 meters.
Maximum frequency, 98 per cent at station 140; other
records above minimum (2 to 78 per cent) from stations 15,
17, 18, 19, 20, 21, 22, 23, 24, 25, 29, 33, 38, 46, 67, 69, 70, 81,
85, 95, 99, 109, 131, 137, 14s, 148, 150. 151, 152, 155, 156,
157, 158, 159, 160; averages in net samples, 7.1 and 17. 1 per
cent in the Atlantic and Pacific, respectively; in pump sam-
ples there were 1 to 12 loricae; average in the Pacific, 2.

Temperature: Atlantic, pumpsamples i4?6o-27?88 (22^23),
net samples i8?o8~37?i8 (24?go); Pacific, i4?77-29?2i
(24?4o) and u?48-28?74 (23?7o), respectively. Salinity:
Atlantic, pump samples 35.61-37. is (36.00), net samples
35.22-37.15 (36.26); Pacific, 33.68-36.46 (35.01) and 3 1 .68—
36.42 (34.72), respectively. Density: Atlantic, pump samples
23.26-26.62 (25.65), net samples 22.84-26.07 (24.81); Pacific,
21.60-25.91 (23.99) and 20.20-26.50 (23.60), respectively.
pH: Atlantic, pump samples 7.93-8.28 (8.18), net samples
8.09-8.32 (8.24); Pacific, 7.86-8.47 (8.28) and 7.68-8.39
(8.23), respectively.

EPIORELLA Kofoid and Campbell

Epiorclla Kofoid and Campbell, 1939, p. 134.

Epiorella includes species which were formerly included
in Epiplocylis, but which have a collar. Kofoid and Camp-
bell (1939) define these as a new genus. Epiorella evidently
arose from Epiplocylis by the addition of the collar, as
Codonaria arose from Codonclla.

TAXONOMY AND DISTRIBUTION

73

Epiorella frequently occurs in cooler waters than does
Epiplocylis, often, in the tropics, being in deep water at
lower than surface temperatures. It frequently occurs in
dense swarms where nearly all the Tintinnoina are oi one
species.

Four species arc described here.

Epiorella acuta Kofoid and Campbell
Epiplocylis acuta Kofoid and Campbell. 1929, pp. 175-176, fig.
3---

The short, distinctly subcorneal lorica, with free lines on
the upper half, has a length of 1.6 oral diameters. The low
sudoral cuff surrounds the oral rim. The cup flares (87 )
within the anterior 0.1 oral diameter, and its free edge forms
a ledge with a diameter of 1.22 oral diameters; a trough
separates cuff and ledge. The cup tapers (7 ) in the upper
half ami then becomes convex conical (65 ); the diameter
at the level of change is 0.89 oral diameter. The aboral
horn is short (0.22 oral diameter), conical (25 ), and
sharply pointed.

The thin wall (0.02 oral diameter) has laminae with en-
closed small alveoles. The cuff is hyaline, the ledge and cup
denser. The upper bowl (0.45 total length) has 64 free
lines, which are commonly interrupted, anastomosed, and
generally vertical. The lower section has large pentagonal
areas with heavy walls and sunken enclosures.

Length, 72U.

Epiorelln acuta resembles most closely E. ralumensis, but
it has a region of free lines well differentiated, whereas
ralumensis has a fully reticulated surface. The aboral end
and horn are longer in ralumensis. Epiorella curta is also
conical, but has scarcely any horn.

Recorded from nineteen stations in the Pacific, as follows:
two (36, 38) in the Pacific equatorial region, thirteen (40,

41, 42, 43, 44, 45, 46, 69, 70, 71, 72, 74, 77) in the Galapagos
region, two (49, 85) in the region of South Pacific island
fields, one (108) in the North Pacific trade region, and one
( 1 1 3 ) in the North Pacific middle latitudes.

There are 22 pump and 15 net samples, of which 14 were
taken at the surface, 12 at 50 meters, and 11 at 100 meters.
Maximum frequency, 54 per cent at station 70; other records
above minimum (2 to 10 per cent) from stations 38, 40, 41,

42, 69, 71, 74, 113; average in net samples, 8.9 per cent; in
pump samples there were 1 to 128 loricae, average 9.8.

Temperature: pump samples i2?73~26?54 (i9?8o), net
samples i4?55~27?94 (20?24). Salinity: pump samples
31.62-36.17 (35-49). net samples 32.88-36.25 (34.99). Den-
sity: pump samples 20.34-26.28 (23.08), net samples 21.31-
26.28 (24.63). pH: pump samples 7.85-8.25 (8.04), net
samples 7.68-8.89 (8.10).

Epiorella brandti Kofoid and Campbell

Epiplocylis brandti Kofoid and Campbell, 1929, p. 177, fig. 324.

The short, rather stout, wide lorica, with free lines limited
to the upper third, has a length of 1.18 oral diameters. The
erect suboral cuff surrounds the thin oral rim. The cup

flares (70°) with a narrow ledge the diameter of which is
1 . 1 1 oral diameters. It is separated from the cuff by a
shallow trough. The lower part of the cup is convex conical
(18 in the anterior half and increasing from 6o° to ioo° in
the lower section)- The aboral horn is short (0.1 oral
diameter) and conical (18 ), and its free tip is sharp.

The suboral cuff is finely reticulated with minute primary
alveoles, the ledge is dense, and the cup proper translucent.
The wall is 0.02 oral diameter in thickness across the cup.
The free lines are about 30 in number, but some anastomose
and all are more or less crooked in course. The lower two-
thirds of the cup is reticulated, as is the horn, with large,
prevailingly pentagonal meshes, the walls oi which arc raised
above the enclosed material. The horn, as in other species,
is solid.

Length, 59 to 70^.

Epiorella brandti has free lines and a less saccular bowl
than E. reticulata. Its bowl is not so definitely conical nor
its horn so long as in E. acuta or E. ralumensis. None of the
species in this collection resembles E. frcymadli of the Indian
Ocean.

Recorded from twelve stations in the Atlantic, as follows:
two (2, 16) in the Gulf Stream, two (17, 18) in the Sargasso
Sea, six (22, 23, 24, 25, 26, 27) in the Atlantic equatorial
region, and two (31, 33) in the Caribbean Sea.

There are 5 pump and 13 net samples, of which 1 was
taken at the surface, 9 at 50 meters, and 8 at 100 meters.
Frequency, 2 per cent at stations 2, 17, 18, 26, 27; other
records all minimum; averages, 1.2 loricae and 1.6 per cent
in pump and net samples, respectively.

Temperature: pump samples, 23?i2-28?25 (25?73), net
samples i4?6o-26?04 (20^73). Salinity: pump samples
35.90-36.25 (36.09), net samples 35.70-36.81 (36.39). Den-
sity: pump samples 23.23-24.67 (23.94), net samples 23.98-
26.07 (25.55). pH: pump samples 8.14-8.30 (8.22), net
samples 7.93-8.30 (8.26).

Epiorella curta Kofoid and Campbell
Epiplocylis carta Kofoid and Campbell, 1929, p. 178, fig. 319.

The short, saccular lorica, with clear-cut free lines on the
upper half, has a length of 1.73 oral diameters. The thin
oral margin is surrounded by the low, incurved suboral cuff.
The cup or bowl flares (30 ) within the anterior 0.14 oral
diameter, and its diameter at the free end is nearly 1.2 oral
diameters. It is separated from the cuff by a shallow trough.
The section below has a length of 0.91 oral diameter and its
diameter at the lower end is 1.09 oral diameters. The lower
end of the bowl is subhemispherical (87 ), with very full,
distinctly convex sides. The short (0.18 oral diameter),
conical (40°) aboral horn is sharply pointed.

The cuff is hyaline, and the bowl, especially around the
flaring region, is denser. The wall has a thickness of 0.02
oral diameter or less. The upper half has about 40 longi-
tudinal, commonly anastomosing free lines, and the lower
bowl has an irregular meshwork of prevailingly pentagonal
deep areas, as does also the horn.

Length, 761.1.

74

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

The Carnegie loricae are more saccular aborally than is
usual for the species.

Epiorella curta is more elongate and has less regular free
lines than E. brandti. The free lines are longer than those
of E. reticulata, and the aboral region is more conical than
in E. healdi.

Recorded from two stations (41, 42) in the Galapagos
region.

There are 3 net samples, 1 taken at the surface and 2 at
100 meters. Frequency, 22 per cent at station 41 at the
surface; 8 per cent at station 4r, and 18 per cent at station
42, both at 100 meters; average, 16 per cent.

Temperature, I4?33~20?42 (i6?43); salinity, 34.19-35.04
(34-75); density, 24.06-26.17 (25.44); P H , 7.91-8. n (7.98).

Epiorella healdi Rofoid and Campbell

(Figure 79)

Epiplocylis healdi Kofoid and Campbell, 1929, p. 1S0, fig. 321;
Marshall, 1934, p. 643, fig. 16.

The small, cup-shaped lorica, with many free lines on the
upper two-thirds of the bowl, has a length of 1.5 oral diam-
eters. The low, faintly crenulated suborai cuff is a low
cylinder and surrounds the oral margin. The cuff is sepa-
rated from the cup below by a shallow trough. The cup or
bowl proper flares (50 ) for a distance of about 0.13 oral
diameter, and its free margin is 1.13 oral diameters in diam-
eter. The lower part tapers (19 ) for a distance of approxi-
mately 0.66 total length, the diameter at the lower end being
nearly 0.83 oral diameter. The posterior section is narrow
oval, increasing from 42 in its upper part to ioo° in the
lower. At the aboral end is a short (0.2 oral diameter),
conical (17°) horn with a blunt free tip.

The wall is thin, hardly exceeding 0.02 oral diameter in
the cup. The cuff is hyaline and the bowl only a little
denser. The upper two-thirds of the bowl has subvertical,
frequently anastomosing free lines, which reach right to the
free edge of the suborai flare. The posterior third is heavily
reticulated with large, deep pentagonal to hexagonal areas,
as is also the horn.

Length, yi[l.

The Carnegie specimens have more free lines (22 to 28)
than the loricae recorded by Marshall (1934), and conform
to type in shape more closely than these last. The one
figured here (fig. 79) is typical of the species. Perhaps
Marshall's are of another, undifferentiated species.

Epiorella healdi has long free lines on the upper bowl,
unlike E. reticulata. There is in this respect no intergrada-
tion. The lorica is also less wide and less full aborally. Its
aboral end is less conical than that of E. curta, but not so
sharply pointed as in E. ralumensis or E. acuta. Epiorella
brandti is generally more conical and has much shorter free
lines.

Recorded from seven stations (40, 41, 42, 43, 71, 72, 73)
in the Galapagos region.

There are 10 pump and 4 net samples, of which 5 were
taken at the surface, 5 at 50 meters, and 4 at 100 meters.

Maximum frequency, 53 per cent at station 40; other records
above minimum (2 to 29 per cent) from stations 41, 42, 43,
71, 72, 73; averages, 4.6 loricae and 31.7 per cent in pump
and net samples, respectively.

Temperature: pump samples i4?55~25?27 (i9?65), net
samples i4?33~2o?92 (i6?28). Salinity: pump samples 34.80-
35-4 2 (35-I7), net samples 34.19-35.04 (34.78). Density:
pump samples 23.60-26.11 (24.95), net samples 24.06-26.51
(25.54). pH: pump samples 7.92-8.21 (8.05), net samples
7.91-8.11 (7.95).

EPICANCELLA Kofoid and Campbell
Epicancclla Kofoid and Campbell, 1929, p. 173.

Epicancella has dominant vertical and distinct horizontal
bars and ribs, the horizontal elements being supplementary
to the vertical structures. Relationship to Rhahdonella on
the one hand, and to Epiplocylis on the other, is indicated;
the genus may have arisen from the latter.

Epicancella occurs in warm, tropical oceans.

There is but one species.

Epicancella nervosa (Cleve) Kofoid and Campbell

Epicancclla nervosa, Kofoid and Campbell, 1929, p. 173, fig.
318.

The transparent, conical lorica has a recurved lip, and
raised lattice work on the surface; its length is 1.6 oral diam-
eters. The oral margin is thin and the hyaline collar below
it spreads from its base, forming a cuff with a width of con-
siderably less than 0.1 oral diameter. The conical bowl
forms a flattened shoulder below the cuff, which shoulder
rounds over and decreases regularly in diameter; its diam-
eter, at the widest part of the shoulder, is 1.17 oral diameters.
The bowl is a narrow cone (6°) in the anterior half, and
then a wider one (52 ) in the lower section. Its sides are
full, and the bowl as a whole is decidedly convex. The
aboral end is pointed but not prolonged or otherwise
modified.

The surface of the lorica is strikingly unique. The cuff
is glass-clear, and the shoulder region is alveolar with more
or less rounded pentagons of mixed sizes. The remaining
part of the bowl has 16 to 18 subvertical or slightly left-
wound ribs, possibly correlated in their number with the
number of membranelles. These ribs are ridges, and they
branch horizontally so that each interface between two ribs
has 14 to 17 crossbars. These crossbars may rarely branch
again with small vertical ridges, especially near the equa-
torial region of the bowl. The crossbars and their ridges are
never so conspicuous as the vertical ribs. The whole lorica
is almost transparent, and not easy to see in a brightly illu-
minated microfield save as the ribs are darker than the
general outline of the bowl; the denser shoulder region is in
contrast with the rest of the bowl. The cavity follows the
outer contour, and the wall is exceedingly thin.

Length, 80 to 82^1.

Epicancella nervosa is so definitely unique among the Tin-

TAXONOMY AND DISTRIBUTION

75

tinnoina that confusion with other species is unlikely. The
dominant vertical ribs suggest relationship with Rhabdonella,
hut horizontal crossbars are not developed in that genus,
although anastomoses of vertical ribs suggest a tendency
toward their definite differentiation. From any species of
Epiplocylis, E. nervosa differs in that the vertical ribs extend
the whole length of the bowl, and none of the members of
that genus have crossbars.

Recorded from thirty stations, seven in the Atlantic and
twenty-three in the Pacific, as follows: five (17, 18, 19, 20,
21) in the Sargasso Sea, two (28, 29) in the Atlantic equa-
torial region, one (45) in the Galapagos region, four (52,
54, 63, 64) in the South Pacific middle latitudes, two (83,
84) in the region of South Pacific island fields, eight (133,
134, 135, 136, 137, 146, 147, 148) in the California region,
four (138, 139, 140, 150) in the North Pacific trade region,
and four (141, 142, 144, 145) in the North Pacific middle
latitudes.

There are 29 pump and 12 net samples, of which 3 were
taken at the surface, 14 at 50 meters, 23 at 100 meters, and
1 at 150 meters. Maximum frequency, 8 per cent at stations
136, 144; other records above minimum (2 to 6 per cent)
from stations 18, 19, 20, 21, 54, 64, 133, 135, 141, 142, 145,
147, 148; averages, 2.6 and 2.1 loricae in Atlantic and Pacific
pump samples, and 1.7 and 2.6 per cent in Atlantic and
Pacific net samples, respectively.

Temperature: Atlantic, pump samples 2o?32-27? 11 (22?78),
net samples 20?32-26?79 (23?i5); Pacific, i4?73~27?46
(i8?94) and i8?74~20?o7 (i9?3i), respectively. Salinity:
Atlantic, pump samples 36.22-36.81 (36.59), net samples
36.31-37.15 (36.73); Pacific, 34.27-36.49 (34.96) and 34.32-
35.40 (34.90), respectively. Density: Atlantic, pump samples
23.62-26.07 (25.18), net samples 23.79-26.07 (25.20); Pacific,
22.90-25.48 (24.53) and 24.24-25.37 (24.88), respectively.
pH: Atlantic, pump samples 8.20-8.29 (8.23), net samples
8.19-8.27 (8.24); Pacific, 8.07-8.39 (8.26) and 8.16-8.39
(8.27), respectively.

XYSTONELLIDAE Kofoid and Campbell

Xystonellidae Kofoid and Campbell, 1929, p. 225.

Included in this family are four genera: Parafavella,
Xystonellopsis, Parundclla, and Xystonella. All four arc
present in the material of this expedition. The family is
widely spread in most seas except the Antarctic.

PARAFAVELLA Kofoid and Campbell

Parafavella Kofoid and Campbell, 1929, pp. 157-158.

Parafavella is one of the most interesting genera of the
Tintinnoina in extent of speciation and extraordinary num-
bers of individuals; the latter fact offers extensive oppor-
tunity for statistical investigation on a quantitative basis,
which should at some time be undertaken in special memoirs.
The geographical distribution of the genus is no less inter-
esting. Limited to regions of low temperature and never ex-

tending far below about latitude 45 north, Parafavella offers
opportunity for study of the effects of latitude and tempera-
ture on evolution. Certain species are limited to the Pacific,
and others to the North Atlantic. Only battered specimens
extend the range south of the arctic regions, where the genus
reaches its greatest development in numbers and kinds;
perhaps polar seas will reveal yet more when these are more
closely examined.

Eleven species are described here, of which one is new, and
another is designated by a new name.

Parafavella affinis, new species
(Plate 1, figure 5)

The lorica is bell-shaped, rather stout, and 2.05 oral diam-
eters in length. The oral margin carries a crown of about
26 sharp, spikclike teeth; these spread outward at about 6o°
from the horizontal, and are evenly spaced, narrow, and
rounded in section. The conical bowl (30 ) may be sub-
divided into two regions. The anterior section has a length
of 1.1 oral diameters; it is an inverted truncated cone (12 ),
and the diameter at the aboral, truncated end is 0.74 oral
diameter; the sides are full, distinct arcs. The aboral sub-
division has as its base the lower end of the section above it;
it is conical (45°), with full sides, and is 0.85 oral diameter
in length. The aboral end is the simple, sharp end of this
cone, and lacks special differentiation.

The wall is evenly and smoothly contoured, with not the
slightest trace of irregularity. It is uniformly less than 0.03
oral diameter in thickness. It has, in section, typical second-
ary prisms in a single layer enclosed by laminae. The
surface has subuniform hexagons at all levels, except near
the aboral tip and in the teeth. The cavity follows the con-
tour exactly.

Length, 1071.1; oral diameter, 52U..

Parafavella affinis resembles P. pacifica closely in general
form, but has a less elongated upper bowl and fewer teeth
(26 as against 32 to 42). It bears some likeness to P. acumi-
nata, but its margins bear teeth, and there is a less sharp
distinction between upper and lower bowls. Parafavella
acuminata is figured by Kofoid and Campbell (1929) as
P. greenlandica.

Recorded from ten adjacent stations in the Pacific, as fol-
lows: two (117, 126) in the North Pacific middle latitudes,
six (118, 119, 120, 121, 122, 123) in the East Asiatic mar-
ginal sea, and two (124, 125) in the Alaskan secondary
region.

There are 25 pump and 14 net samples, of which 13 were
taken at the surface, 14 at 50 meters, and 12 at 100 meters.
Parafavella affinis occurs equally at all levels examined.
Frequency, 80 per cent at station 124, at the surface; 500
loricae counted in the pump sample at station 119, at 50
meters; all records, except that at station 118, above mini-
mum (2 to 28 per cent); average in net samples, 22.7 per
cent; in pump samples, 52 loricae (1 to 500).

Temperature: pump samples 2?oi-i2?56 (6?2i), net
samples 2?oi-8?22 (5?io). Salinity: pump samples 32.63-

7 6

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

34.22 (33-03), net samples 32.70-33.69 (33.14). Density:
pump samples 24.90-26.52 (25.44), net samples 25.74-26.76
(26.09). pH: pump samples 7.85-8.21 (7.86), net samples
7.64-8.03 (7.70).

Type locality, station 121, at 50 meters; latitude 46 05'
north, longitude 171 ° 32' east.

Parafavella cylindrica (Jorgensen) Kofoid and Campbell

(Figure 51 )
Parafavella cylindrica, Kofoid and Campbell, 1929, p. 162, fig.
312-

The tall, narrow, elongated lorica, with strong teeth, taper-
ing bowl, and rounded aboral end, has a length of 5.8 oral
diameters. The oral margin has 24 sharply pointed, narrow,
triangular teeth which are curved outward and the length of
which is less than 0.1 oral diameter. These teeth rest on a
very low, conical (54 ) crown or collar, the length of which
is 0.08 oral diameter, the diameter being 0.93 oral diameter
at the lower end. Below this contracted neck the inverted,
conical (2 ), decidedly elongated bowl arises. This section
has a diameter of 1.06 oral diameters just below the neck,
and 0.84 at its bottom end, which is 0.84 total length below
the oral rim. The bowl contracts below this last level in the
shape of an approximate half-ellipse, being about 45 in its
anterior half and increasing to 8o° in the posterior section.
Its length is 0.67 of its own diameter, and its sides are con-
vex. At the aboral end is a short, curved, minutely blunted,
conical (15°) aboral horn, the length of which is about 0.37
oral diameter.

The wall has a maximum thickness of 0.06 oral diameter
near the upper end of the tapering bowl, and gradually this
lessens to as little as three-tenths as much. Thin laminae
enclose a single layer of radial, rectangular prisms. There
is a meshwork of practically uniform, double-walled hexa-
gons which enclose hyaline contents. The aboral horn is
hollow and the cavity follows the outer contour.

Length, 363^.

The Carnegie loricae have fewer, longer teeth than usual,
as well as slight suboral contraction and more taper.

Parafavella cylindrica compares nicely with P. subrotun-
data, but is longer and more slender, with an elliptical rather
than rounded aboral end, and without a sharp, pointed horn.
It is longer than P. dilatata, much less tapering, with fewer
teeth, and with a somewhat different aboral region. Para-
favella gigantea has a longer horn and more teeth, and
P. r obits ta has a longer horn and shorter bowl.

Recorded from five stations, one in the Atlantic and four
in the Pacific, as follows: one (6-7) in the North Sea, and
four (118, 120, 121, 122) in the East Asiatic marginal sea.

There are 9 net samples, of which 3 were taken at the
surface, 4 at 50 meters, and 2 at 100 meters. Frequency, 96
per cent at station 118 at 100 meters; all other records, save
that in the Atlantic, above minimum (2 to 70 per cent);
average in the Pacific, 27.1 per cent.

The following data apply to the Pacific stations only:
temperature, 2?oi-io?i8 (5?52); salinity, 32.06-33.78
(33.29); density, 25.14-26.57 (26.23); pH, 7.86-8.21 (7.98).

Parafavella denticulata (Ehrenberg) Kofoid and Campbell
(Figure 50)

Parafavella denticulata, Kofoid and Campbell, 1929. p. 163, fig.

310; Hada, 19326, pp. 564-565.
Parafavella dentiula (lapsus) Hada, 1932a, pp. 50-51, fig. 15.
Vavella denticulata, Schulz and WulfT, 1929, pp. 344-352, figs.

2-25, pis. 12, 13; also var. tenuis forma arctica, fig. zie-g.

The tall lorica, with stout teeth, subcylindrical bowl, con-
tracted aboral region, and moderately long aboral horn, has
a length of 3.76 oral diameters. The thin oral rim has 32
short, relatively wide, subequidistant, uniform, suberect,
triangular teeth. The slightly tapering (2 ) anterior bowl
occupies about 0.55 total length, and its diameter at the lower
end is near 0.93 oral diameter; its walls are without traces of
minute irregularity. Below its lower end the bowl gradually
contracts in the form of a half-ellipse, the length of which is
1. 17 its own diameter. It is at first 30 and increases to 52
in the posterior three-tenths. The aboral horn is conical
(15°) and its length reaches 0.72 oral diameter. Its free tip
is minutely blunted.

The wall has a uniform thickness of nearly 0.03 oral diam-
eter except in the horn, where it is somewhat more. There
are thin laminae and enclosed radial, rectangular prisms.
The outer surface has distinct, although faint, hexagonal
prisms of modest size. The cavity approximates the outer
contour.

Length, 253^.

The Carnegie loricae lack the suboral flare evident in
typical loricae. Schulz and WulfT (1929) illustrate and dis-
cuss a wide variety of forms, only some of which belong to
denticulata (sensu stricto), the others belonging to several
distinct forms, some of which may be new. The variability
of Parafavella is certainly very great, and physical influences
profoundly affect formation. The effects of temperature,
salinity, density, and hydrogen-ion concentration are yet to
be experimentally studied in these ciliates. Observational
data are suggestive.

Parafavella denticulata resembles P. gigantea, but is shorter
and stouter, with a less lengthy horn. It is not so stout as
P. robusta, and there is a sharper transition in lower bowl
and horn than in P. obtusangula. It has numerous teeth,
lacking in the somewhat similar P. edentata. The other
species are so distinct that they could scarcely be confused
with denticulata.

Recorded from seven stations, three in the Atlantic and
four in the Pacific, as follows: two (between stations 6 and
7) in the North Sea, one (13) in the American cold-water
region, two (116, 117) in the North Pacific middle latitudes,
one (118) in the East Asiatic marginal sea, and one (130)
in the California region.

There are 3 pump and 10 net samples, of which 5 were
taken at the surface, 6 at 50 meters, and 2 at 100 meters.
Maximum frequency, 95 per cent at station 6-70; other
records above minimum (2 to 56 per cent) from stations 116,
117, 118, 130; average in Pacific net samples, 12. 1 per cent.

Temperature: Atlantic, pump sample n ^27, net sample
i?64; Pacific, 8?33 and 8?93~i6?07 (11^54), respectively.

TAXONOMY AND DISTRIBUTION

77

Salinity: Atlantic, pump sample 32.68, net sample 33.40;
Pacific, 33.72 and 33.06-34.22 (33.83), respectively. Density:
Atlantic, pump sample 24.94, net sample 26.90; Pacific, 26.24
and 24.99-26.41 (25.75), respectively, pll: Atlantic, not
recorded; Pacific, pump sample 8.21, net samples 7.98-8.26
(8.12). The data for this species, as for others taken from
intermediate stations, are incomplete.

Parafavella dilatata (Jorgensen) Kofoid and Campbell
Parafavella dilatata, Kofoid and Campbell, 1929, pp. 163-164,

fig- 315-
The tall, narrow, hell-shaped lorica. with long bowl and
short horn, has a length of 2.94 oral diameters. The thin
oral margin has approximately 60 subequidistant, subequi-
lateral, triangular teeth, which invariably turn inward. The
contracted conical section of the bowl (12 ) has a length of
0.7 total length, and its diameter at the lower end is 0.53
oral diameter. This section contracts to 0.7 oral diameter at
0.4 total length from the rim. The aboral region of the
bowl is convex conical (52°) and has a length of nearly 0.17
total length. The aboral horn is a narrow, pointed, barely
bulging cone (7 ) with a length of almost 0.14 total length.
The wall is subuniformly 0.05 oral diameter in thickness,
with little trace of laminae but with well developed large,
rectangular radial prisms in a single layer. The outer mesh
is made up of uniformly rather small hexagons, except that
the horn is glassy. The lumen follows the outer contour and
continues to the tip of the closed horn.
Length, 247 to 3901.1.

This single lorica resembles Brandt's rather more than it
does the one figured by Jorgensen.

Parajavella dilatata differs from P. subrotundata in having
a more clearly defined campanulate bowl, and it also has sub-
median contraction; this last character serves to distinguish
dilatata from all other species. If it is only a distorted (con-
tracted) P. cylindrica, as Kofoid and Campbell suggested,
then dilatata, cylindrica, and subrotundata are possibly only
extremes of one variable species, the name of which would
have to be subrotundata by plate precedence.

Recorded from one station (6-7) in the North Sea, in a
net sample taken at the surface. Frequency, minimum.

No physical data accompany this and other collections be-
tween the regular series stations.

Parafavella edentata (Brandt) Kofoid and Campbell
Parajavella edentata, Kofoid and Campbell, 1929, p. 164, fig.

296.
The rather short, campanulate lorica lacks oral teeth but
has a relatively long aboral horn. Its length is 2.2 oral diam-
eters. The oral margin is the simple, unmodified upper edge
of the bowl. The bowl is virtually a cylinder in the upper
two-thirds, with a little suboral flare in some loricae and
slight local inflation and contraction in others, and with the
diameter at the lower end of the cylinder 0.83 to 1.00 oral
diameter. The aboral section is about 0.75 oral diameter in
length; it is a plane -to-concave cone (55°). The aboral horn
is 0.5 oral diameter in length, narrow conical (8°), and
sharply pointed.

The wall is nearly 0.04 oral diameter in thickness, more
or less subuniformly. There are exceedingly thin laminae,
between which is a single layer of radial prisms. The outer
surface has a distinct hexagonal meshwork of small suboral
prisms and larger median ones, and is entirely devoid oi
prisms in the greater part of the horn. The horn is hollow.

Length, 80 to 150U.

There is marked diversity in length and proportions and
also in the relative length of the howl, which varies from
short to long.

Parajavella edentata resembles P. subedentata, but is
usually relatively stouter and more widely campanulate, and
has a more gradual transition between bowl and horn. The
remaining edentulous species, P. digitalis, P. inflata, and P.
curvata, lack aboral horns entirely or else have only points,
have long cylindrical bowls, and are longer. Parajavella
denticulata resembles edentata, but always has strong teeth;
however, edentata is not an edentate form of that species.
Recorded from one station (7) in the North Sea, in a net
sample taken at the surface. Frequency, minimum.

Temperature, 8^92; salinity, 35.21; density, 27.31; pH,
8.08.

Parafavella gigantea (Brandt) emended Kotoid
and Campbell
(Figures 52, 53)
Parajavella gigantea, Kofoid and Campbell, 1929, p. 165, rig.

311; Hada, 1932a, pp. 51-52, fig. 16; 1932/;, p. 565.
The large, tall, generally cylindrical lorica, with narrow,
contracted conical aboral region and very narrow, pointed
aboral horn of moderate length, has a length of 5.9 to 6.9
oral diameters. The oral margin has approximately 46
equidistant, narrow triangular, outward-flaring (45°) teeth,
nearly 0.1 oral diameter in length. The long bowl is cylin-
drical to barely tapering (least diameter 0.77 oral diameter
near 0.7 total length), and is without local modification in
contour. The lower, shorter, slightly convex conical (40 to
45°) part has a length of barely 1.0 oral diameter. The
aboral horn is narrow conical (5 to 16 ), pointed at the
free tip, and sometimes as much as 1.17 oral diameters in
length.

The wall is slightly thickened suborally but averages 0.06
oral diameter. It has very distinct rectangular radial prisms,
and the surface meshwork is of uniform small hexagons in
well developed pattern save in the horn.

Length. 337 to 57 6 l l -

The Carnegie loricae lack contraction in the bowl, being

either directly cylindrical or tapering, and are much more
like that figured by Kofoid and Campbell than that figured
by Hada.

Parajavella gigantea is longer than P. denticulata and has
a longer horn, but not so long as that of P. promissa, which
species is also shorter. Parajavella gigantea differs also from
P. cylindrica in horn, the latter having only a short stub.
Parajavella acuta is shorter and has an acuminate aboral end.
The bowl of P. gigantea is longer than that of P. elegans,
which species is shorter. It differs from P. robusta in its
more slender contour and in being longer.

7»

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Recorded from eleven stations, five in the Atlantic and six
in the Pacific, as follows: two (between stations 6 and 7) in
the North Sea, one (9) in the Atlantic drift, two (12, 13) in
the American cold-water region, two (116, 117) in the North
Pacific middle latitudes, three (118, 122, 123) in the East
Asiatic marginal sea, and one (130) in the California region.

There are 7 pump and 13 net samples, of which 8 were
taken at the surface, 5 at 50 meters, and 7 at 100 meters.
Maximum frequency, 97 per cent at station 122; other
records above minimum (2 to 91 per cent) from stations
6-ja, 6-yb, 9, 13, 117, 118, 122, 130; averages, 11.4 and 55.4
per cent in Atlantic and Pacific net samples, respectively; in
Atlantic pump samples, 9.8 loricae.

Temperature: Atlantic, net samples i?64-ii?27 (8?n);
Pacific, pump samples 2?54-i8?22 (6°gy), net samples 2°^-
io?i8 (6?9o). Salinity: Atlantic, net samples 32.68-35.14
(33.71); Pacific, pump samples 32.84-34.06 (33.01), net
samples 32.76-33.78 (33.27). Density: Atlantic, net samples
24.94-26.90 (26.22); Pacific, pump samples 25.58-26.45
(26.20), net samples 25.52-26.57 (26.05). pH: Atlantic, net
samples 8.08-8.10 (8.09); Pacific, pump samples 7.90-8.02
(8.00), net samples 7.90-8.21 (8.00).

Parafavella hadai, new name

Not Favella denticalata forma subcylindrica Schulz and Wulff,

1929, p. 345, fig. 21<f.
Parafavella subcylindrica Hada, 1932a, pp. 54-55. fig- -°-

"Lorica short finger-shaped, 2.7 oral diameters in length;
oral margin denticulate with many small triangular teeth
(about 60); bowl subcylindrical, dilated slightly in the sub-
oral 0.16 of the total length, gradually contracting aborally,
convex conical (8°) in the posterior 0.3 of the lorica; aboral
end nearly hemispherical, somewhat subacute, without an
aboral horn.

"Length, 169^1; oral diameter 63(1.

"Differs from Parafavella digitalis Kofoid and Campbell
in having the denticulate oral rim, from P. hemispherica
(Meunier) in numerous oral teeth and in the rounded aboral
end, and from P. rotundata (Jorgensen) in smaller size and
in stouter proportions."

Parafavella hadai occurs at only one station, off Kam-
chatka, recorded by Hada in 1932.

Parafavella hadai is renamed in this report because the
specific name subcylindrica had already been applied by
Schulz and Wulff to another species, which Hada apparently
overlooked. The specific name hadai is, thus, appropriate.
It was not recorded in the material of the Carnegie, although
many of the Japanese species have been reported again by
this expedition.

Parafavella obtusangula (Ostenfeld) emended Kofoid
and Campbell

Parafavella obtusangula, Kofoid and Campbell, 1929, p. 168,

fig. 309.
Favella denticulata var. tenuis, Schulz and Wulff (part), 1929,

p. 344, fig. zog (see also P. denticulata).

The lorica is short and bell-shaped, with an angular lower

bowl, a pointed aboral horn, and a length of 2.8 oral diam-
eters. The oral margin is regularly denticulate with approxi-
mately 36 short, erect, wide triangular teeth. The bowl is
cylindrical in the upper 1.5 oral diameters, and obtusely
conical (46 ) in the lower section, having there plane-to-
barely-convex sides. The aboral horn is short (0.2 oral
diameter) and narrow conical (8°), and has a sharp point.

The wall is within 0.05 oral diameter in thickness uni-
formly at all levels, and has thin laminae which enclose a
single layer of radial, rectangular secondary prisms. The
outer surface has moderately large, rounded hexagons except
in the aboral horn, which is usually hyaline.

Length, 140U.

Schulz and Wulff figure one lorica (fig. log) which be-
longs to this species; the others of their series do not appear
to be related.

Parafavella obtusangula differs from its close relative P.
denticulata in the obtuse, long, conical lower bowl and
shorter horn. The transition between bowi and horn is also
less marked and the cylindrical part of the bowl is shorter.
It is unlike P. parumdentata, which has suboral flare and
much less transition between bowl and horn.

Recorded from one station (117) in the North Pacific
middle latitudes, in a net sample taken at 50 meters. Fre-
quency, 2 per cent.

Temperature, I2?56; salinity, 34.22; density, 25.89; pH,
8.06.

Parafavella parumdentata (Brandt) Kofoid and Campbell

(Figures 49, 54, 55)
Parafavella parumdentata, Kofoid and Campbell, 1929, p. 168,
fig. 306; Hada, 1932a, pp. 49-50, fig. 14.

The lorica is goblet-shaped, small, with evenly contracted
bowl and pointed aboral end lacking a distinct horn, and is
1.9 to 2.1 oral diameters in length. The oral margin has
about 32 incurved, short, equidistant, clawlike, triangular
teeth (omitted in the drawings). The suboral region of the
bowl flares as a low, basal, inverted segment of a cone (46
to 6o°) with a length of within 0.17 oral diameter and with
a diameter at the lower end of within 0.86 oral diameter.
Below this throat the bowl contracts as an inverted cone
(5 ) for about 0.5 total length and then (32° to 48 ) be-
comes a wider cone with full sides. The aboral end is not
prolonged as a pedicel, being but the simple sharp end of the
lower bowl.

The wall is uniformly approximately 0.05 oral diameter in
thickness, but its structure was indeterminate in the loricae
of this expedition. The outer surface has distinct, very small,
elongated hexagons.

Length, 97 to 107U.

The Carnegie loricae are distinctly shorter and stouter than
those included by Kofoid and Campbell, which are 127 to
20o[i and 2.8 to 3.0 oral diameters. The shape of their
figured lorica is somewhat different, as is also that of Hada.
One of the loricae figured here (fig. 49) is aberrant in shape,
proportions, and size.

Parafavella parumdentata, though showing considerable

TAXONOMY AND DISTRIBUTION

79

variation, may be compared with P. pacifica, from which it
differs in the more tapering aboral end and lack of horn;
from P. obtusangula it differs in aboral characters.

Recorded from seven adjacent stations in the Atlantic, as
follows: one (7) in the North Sea, four (8, 9, 10, 11) in the
Atlantic drift, and two (12, 13) in the American cold-water
region.

There are 3 pump and 17 net samples, of which 7 were
taken at the surface, 8 at 50 meters, and 5 at 100 meters.
Maximum frequency, 100 per cent at station 8 at 100 meters,
station 1 1 at the surface, station 12 at 100 meters, and station
13 at 50 meters; all other records 2 to 99 per cent.

Temperature: pump samples 7?oi-l4?47 {\l°.7.j), net
samples i?64-n?27 (7?9i). Salinity: pump samples 32.68-
34.97 (34.18), net samples 33.40-35.25 (34.82). Density:
pump samples 24.94-27.42 (26.38), net samples 24.94-27.96
(27.14). pH: pump samples 7.92-8.06 (7.99), net samples
7.93-8.08 (7.98).

Parafavella promissa Hada
Parafavella promissa Hada, 1932(7, p. 53, fig. 18.
The elongated, narrow lorica, with sharply recurved oral
teeth, cylindrical bowl, and very long, spinelike aboral horn,
has a length of 7.3 oral diameters. The oral margin flares
very slightly, and there are 24 subequidistant, narrow, de-
cidedly outward-flaring, triangular teeth. The long upper
bowl is a cylinder expanded a bit near the upper end, and
occupies 2.56 oral diameters. The aboral convex-conical
(45°) section has a length of nearly 1.0 oral diameter. The
very elongated aboral horn is conical (not over 2°) and is
about 0.5 or more of the total length. At its free tip there
is sometimes a minute disk-shaped expansion.

The wall has a thickness of 0.06 oral diameter in the bowl,
a little more suborally, and somewhat less near the horn.
There are exceedingly thin inner and outer laminae enclosing
a single layer of radially arranged secondary prisms. The
rather small surface meshwork of hexagons covers the bowl
everywhere, but the long aboral horn and the teeth are free
of prisms. The lumen enters the horn and elsewhere pre-
cisely conforms to the outer contour.
Length, 334P.

The specimens of this expedition differ only in minor
ways from Hada's (1932a) figured lorica.

Parafavella promissa bears only slight resemblance to P.
subula. It is longer and much more slender, with a cylin-
drical instead of short, wide bowl, and the stronger, recurved
teeth are prominent. Its bowl and aboral horn are also un-
like those of P. elegans, and it could hardly be confused
with the other species, with which, also, it is seldom asso-
ciated in the Pacific.

Recorded from two stations in the Pacific, as follows: one
(128) in the North Pacific middle latitudes, and one (130)
in the California region.

There are 3 pump and 2 net samples, of which 3 were
taken at 50 meters and 2 at 100 meters. Maximum fre-
quency, 45 per cent at station 130; another record above
minimum (9 per cent) from station 128; averages, 4 and

26.5 per cent in pump and net samples, respectively.
Temperature: pump samples 8?96-i2?9i (u?25), net
samples 8?9&-i2?9i (io?93). Salinity: pump samples 33.07-
337 2 (33-39), net samples 33.40-33.72 (33.56)- Density:
pump samples 25.13-26.14 (25.48), net samples 25.19-26.14
(25.66). pH: pump samples 8.06 8.26 (8.14), net samples
8.06 8.26 (8.16).

Parafavella ventricosa (Jorgensen ) Kofoid and Campbell

Parafavella ventricosa, Kofoid and Campbell, 1929, p. 171, fig.

314; Hada, 1932^, p. 566, fig. iS.
Favella denticulata var. robusta forma arctica Schulz and Wulfl,

1929, p. 345, fig. 2 if.
The rather large, stout, aborally inflated lorica, with a
short aboral horn, has a length of 5.17 oral diameters. The
thin oral margin has about 36 very short, subequidistant,
somewhat curved teeth. The bowl is cylindrical in the an-
terior 0.45 total length and then gradually expands in the
remaining section to 1.25 oral diameter at 0.65 total length
from the rim. Below this level it contracts rapidly as a
convex cone (6o°) with a length of 0.26 total length. At its
lower end is the short (0.4 oral diameter), conical (5°),
pointed aboral horn.

The thin wall (0.05 oral diameter) is subuniform, with
thin laminae and enclosed radial, rectangular prisms in a
single layer. The wall has distinct, rounded hexagons of
nearly uniform size at all levels; the aboral horn is hyaline
near the free tip.
Length, 34611.

Although in some ways differing from the typical form,
the lorica figured by Schulz and Wulff (1929) appears to
belong to this species. The loricae of this expedition con-
form neatly to type.

Parajauella ventricosa differs from all the other species in
the extent of inflation of the aboral region, which has a
decidedly gourdlike shape.

Recorded from one station (121) in the East Asiatic
marginal sea, in a net sample taken at 50 meters. Frequency,
minimum.

Temperature, 3?77; salinity, 33.06; density, 26.28; pH,
7.92.

XYSTONELLOPSIS Jorgensen
Xystonellopsis, Kofoid and Campbell, 1929, pp. 238-239.
Xystonellopsis has very much the same position and
abundance in the tropics as Parafavella in arctic seas. As
regards abundance, it is not so frequent, but is common in
small percentages in nearly any catch in warm seas. In
number of species it ranks somewhat better. The two genera
have in common the simple oral margin, the somewhat
similar form, and close likeness in wall structure. For these
reasons they are allied here. Kofoid and Campbell (1929)
placed Parafavella near Favella, partly for historical reasons,
and because the surface hexagonal patterning of the wall of
Favella, in some species, is much like that in Parafavella.
More careful work and consideration seem to indicate closer
relationship to the Xystonellidae on the part of Parafavella.
Both Xystonellopsis and Parafavella are specialized genera of

8o

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

highly peculiar, special habitats in very different parts of the
ocean. Favella is a more generalized genus of more tem-
perate waters, especially coastal. Few if any species of
Xystonellopsis ever enter neritic environments, this genus
being common in the high seas. It is interesting that none
of the Xystonellidae enter the Antarctic, although the family
has mastered most other seas.

Twenty-two species are described here.

Xystonellopsis abbreviata Kofoid and Campbell

Xystonellopsis abbreviata Kofoid and Campbell, 1929, p. 239,
fig. 468.

The moderately stout lorica, with pulc/ira-\\ke form and
short pedicel, has a length of 5.1 oral diameters. The oral
margin is thin and erect. The suboral band occupies the
anterior 0.16 total length and has 3 rounded, subequal rings
with diameters of nearly 1.27 oral diameters; these rings are
thin, and virtually equidistant, the upper one being a little
below the oral rim. The interspaces between the rings are
thickened. The bowl below the band is subconical (io° in
the upper 0.21 and 38 in the lower 0.16 total length). The
pedicel is thick (0.36 oral diameter at the upper end) and
tapering (5 ), with a slight, aborally truncated skirt at the
distal end. The lance is conical (2 ) and distally pointed,
and has a length of 0.55 oral diameter.

The wall has a thickness of 0.1 oral diameter just below
the spool, and there are thin laminae which enclose small
hexagons in several layers in the spool, radial secondary
prisms in a single layer in the bowl, and large hexagons in
the skirt and lance. The lumen does not enter the spool,
and forms a narrow canal in the lance.

Length, 275 to 336"..

Xystonellopsis abbreviata is much like X. pulchra, but is
not so long; its suboral spool is shorter and the pedicel is
short.

Recorded from one station (108) in the North Pacific trade
region, in a pump sample taken at 100 meters. Frequency,
minimum.

Temperature, 25^36; salinity, 34.99; density, 23.24; pH,
8.23.

Xystonellopsis acuminata Kofoid and Campbell

Xystonellopsis acuminata Kofoid and Campbell, 1929, p. 239,
fig. 482.

The moderately tall lorica, with beros-\ike bowl, stout
pedicel, and pencil-shaped lance, has a length of 5.56 oral
diameters. The oral margin is sharp, with a trace of flare
within the upper tenth. The bowl is subcylindrical in the
anterior half and conical (14°) in the aboral region, which
merges with the pedicel. The lance arises from the barely
differentiated skirt; it is pencil-like, 0.5 oral diameter in
length, and sharply pointed.

The wall is nearly 0.08 oral diameter in thickness, and
the thin laminae enclose several layers of minute alveoles.
On the pedicel are 8 right-twisted (3°) ridges.

Length, 354 to 414ft.

Xystonellopsis acuminata has a clear-cut skirt wholly lack-

ing in X. dahli. It is not so long as X. heros, and has a
pencil-shaped rather than conical lance. Xystonellopsis gaussi
has a stouter, conical lance and more definite pedicel.

Recorded from five stations in the Pacific, as follows: one
(35-36) in the Pacific equatorial region, three (112, 113, 145)
in the North Pacific middle latitudes, and one (131) in the
California region.

There are 1 pump sample and 4 net samples, of which 1
was taken at the surface and 4 at 100 meters. Maximum
frequency, 11 per cent at station 112; other records above
minimum (2 to 8 per cent) from stations 113, 131; average
in net samples, 7.6 per cent.

Temperature: pump sample i6?58, net samples I2?i2-
2I °74 (i8?oo). Salinity: pump sample 34.71, net samples
33-36-347 1 (34-24). Density: pump sample 25.00, net
samples 24.06-25.31 (24.63). pH: pump sample 8.31, net
samples 8.23-8.32 (8.25).

Xystonellopsis armata (Brandt) Kofoid and Campbell

Xystonellopsis armata. Kofoid and Campbell, 1929, p. 241, fig.
481.

The rather tall lorica, with contracted lower bowl, ex-
panded pedicel, and short, conical lance, has a length of 6.65
oral diameters. The oral margin is thin and sharp. The
bowl tapers (5°) in the upper 0.48 total length and becomes
subconical (30 ) in the lower 0.2 total length. The pedicel
increases in diameter from the lower bowl (4°) and has a
length of nearly 0.32 total length; at its distal end is the
squarely truncated, non-expanded skirt. The lance is conical
(32 ), only 0.5 oral diameter in length, and sharply pointed.

The wall reaches nearly 0.1 oral diameter in thickness
suborally, and becomes a little less in the lower bowl. There
are comparatively thick, dark laminae, which enclose homo-
geneous material. The lumen forms a canal in the lance.
The pedicel has 8 subvertical pleats.

Length, 365 to 410U..

Xystonellopsis armata resembles X. gaussi, but the pedicel
becomes distally wider instead of contracted, and the lance
is conical and wide rather than pencil-shaped. It differs
from X. heros in having a subconical lower bowl with dis-
tinct transition to the pedicel, as well as in being shorter. It
lacks the suboral thickening of X. tenuirostris.

Recorded from twenty-one stations, two in the Atlantic
and nineteen in the Pacific, as follows: two (23, 24) in the
Atlantic equatorial region, two (35, 153) in the Pacific equa-
torial region, two (45, 68) in the Galapagos region, three
(65, 66, 67) in the South Pacific middle latitudes, two (109,
150) in the North Pacific trade region, eight (133, 134, 135,
136, 146, 147, 148, 149) in the California region, and two
(142, 145) in the North Pacific middle latitudes.

There are 6 pump and 16 net samples, of which 4 were
taken at 50 meters and 18 at 100 meters. Maximum Ere
quency, 26 per cent at station 67; other records above mini-
mum (2 to 25 per cent) from stations 23, 24, 35, 45, 65, 66,
68, 109, 135, 142, 145, 146, 148, 150; averages in net samples,
1.2 and 11.0 per cent in the Atlantic and Pacific, respectively.

Temperature: Atlantic, net samples 20?99~23?i2 (22?05);

TAXONOMY AND DISTRIBUTION

81

Pacific, pump samples i6?90-28?05 (2i?36), net samples
i4?33-2i?69 (i8?27). Salinity: Atlantic, net samples 36.00-
36.04 (36.02); Pacific, pump samples 34.40-35.02 (34.74).
net samples 34.30-35.21 (34.79). Density: Atlantic, net sam-
ples 24.67-25.30 (24.98); Pacific, pump samples 21.95-25.26
(24.56), net samples 24.24-26.06 (25.04). pH: Atlantic, net
samples 8. 14-8. 18 (8.15); Pacific, pump samples 8.12-8.39
(8.30), net samples 7.88-8.38 (8.18).

Xystonellopsis brandti (Laackmann) Jorgensen
Xystonellopsis brandti, Kofoid and Campbell, [929, p. 241, fig.

474-

The rather short lorica, of extreme narrowness, with sub-
oral swelling and conical pedicel with merged lance, has a
length of 7.8 oral diameters. The oral margin is thin and
erect. The bowl swells in a zone of the uppermost oral
diameter to 1.3 oral diameters at 0.3 oral diameter below the
rim. The upper part of this swollen region has a rounded
shoulder, and the lower part tapers (23 ) and merges with
the bowl below. The bowl below the swollen region swells
to 1.1 oral diameters at 2.8 oral diameters below the rim,
and then is conical ( 12 ) for the remainder of the length of
the lorica. The pedicel-lance occupies about 1.5 oral diam-
eters, and the distal tip is sharp.

The wall has a thickness of nearly 0.2 oral diameter across
the widest part of the bowl, but in the lower bowl is only
0.05 or less. There are thin laminae with faint, fine hexa-
gons in several layers, especially suborally. The lorica is
hyaline, save that the pedicel-lance is dense. There are 6
low, winglike fins which continue from the lower bowl to
the lower pedicel. The lumen is reduced to a canal in the
lance, and does not enter the suboral thickening.

Length, 196 to 22511.

In many respects Xystonellopsis biandti is unique, but it
is possibly somewhat like X. constricts. It is, however,
shorter, lacks the tubular, constricted lower bowl, and has
more suboral thickening.

Recorded from two stations, one each in the Atlantic and
the Pacific, as follows: one ( 19) in the Sargasso Sea, and one
(80) in the Galapagos region.

There are 1 pump and 1 net sample, both taken at 50
meters. Frequency, minimum.

Temperature: Atlantic, net sample 25:31; Pacific, pump
sample 26?o6. Salinity: Atlantic, net sample 37.15; Pacific,
pump sample 35.95. Density: Atlantic, net sample 24.89;
Pacific, pump sample 23.75. pH: Atlantic, net sample 8.27;
Pacific, pump sample 8.19.

Xystonellopsis conicacauda Kofoid and Campbell
Xystonellopsis conicacauda Kofoid and Campbell, 1929, p. 243,

fig. 465.
The fairly short lorica, with duplicated skirts and short,
conical lance, has a length of 5.7 oral diameters. The oral
margin is thin and erect. The bowl swells rapidly to a
diameter of 1.93 oral diameters within the anterior 0.1 oral
diameter, forming a thickened ledge. The lower part of the
swollen region extends for the anterior 0.25 total length.

gradually being reduced posteriorly. Below this region the
bowl becomes subcorneal (io°) for the remainder of its
length. At the lower end it swells to form the upper, ex-
panded skirt. Below this skirt is a short cylindrical section
(approximately 1 oral diameter in length) with a slight skirt
at its lower end. From the center of this skirt arises the
lance, which is conical (6o°), with ils length less than its
basal diameter.

The wall has a thickness of 0.28 oral diameter across the
ledge. The lumen does not enter the ledge, but does follow
the external contour of the lance. The wall is otherwise like
that of X. paradoxa.

Length, 188 to 229a.

Xystonellopsis conicacauda resembles X. paradoxa, but its
lance is shorter, wider, and more conical; the bowl is pro-
portionately longer.

Recorded from one station (19) in the Sargasso Sea, in a
net sample taken at 100 meters. Frequency, minimum.

Temperature, 22?42; salinity, 37.05; density, 25.67; pH,
8.25.

Xystonellopsis crassispinosa Kofoid and Campbell
(Figure 64)
Xystonellopsis crassispinosa Kofoid and Campbell, 1929, p. 244,
fig. 460.

The fairly short lorica, of general cymatica type but with
thick pedicel and stout lance, has a length of 4.7 oral diam-
eters. The oral margin is thin and erect. The bowl swells
immediately below the rim to form a rounded, thick ledge
with a diameter of 1.28 oral diameters; the ledge has a width
of 0.32 oral diameter. The bowl below the ledge is sub-
conical (8° increasing to 33 in the aborai 0.76 oral diam-
eter). The pedicel is thick (0.25 oral diameter) and tubular,
with a length of 0.32 total length, and with a slightly ex-
panded skirt at its distal end. The skirt has 8 (12) clawlike
points, and above each is a short pleat which reaches the
lower end of the pedicel. The lance arises from the center
of the skirt, has a length of 0.6 oral diameter, and is stout,
conical (16 ), and distally blunted.

The wall is thickest suborally, where it reaches nearly 0.09
oral diameter; it tapers to less than three-tenths as much in
the lower bowl. There are thin laminae, which enclose
radial, secondary rectangles, save in the ledge, where there
are several layers of small hexagons. The lumen does not
enter the ledge.

Length, 244 to 265^.

Xystonellopsis crassispinosa differs from X. cymatica and
related species in the stoutness of the pedicel and lance, and
from X. spicata in the narrower ledge.

Recorded from ten stations, three in the Atlantic and
seven in the Pacific, as follows: one (18) in the Sargasso Sea,
two (22, 23) in the Atlantic equatorial region, two (41, 45)
in the Galapagos region, one (95) in the region of South
Pacific island fields, two (136, 146) in the California region,
one (145) in the North Pacific middle latitudes, and one
( 150) in the North Pacific trade region.

There are 1 pump and 9 net samples, of which 1 was
taken at 50 meters and 9 at 100 meters. Maximum frequency.

82

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

20 per cent at station 145; other records above minimum (4
to 6 per cent) from stations 41, 95, 136, 146; average in
Pacific net samples, 6.8 per cent.

Temperature: Atlantic, pump sample 20?32, net samples
20?99-24?44 (22?7i); Pacific, net samples i4?55~28?74
( 1 9°54). Salinity: Atlantic, pump sample 36.81, net samples
36.04-36.18 (36.11); Pacific, net samples 34.32-35.33 (34-53).
Density: Atlantic, pump sample 26.07, net samples 24.42-
25.30 (24.86); Pacific, net samples 22.43-26.11 (24.50). pH:
Atlantic, pump sample 8.21, net samples 8.14-8.21 (8.17);
Pacific, net samples 7.92-8.39 (8.22).

Xystonellopsis cyclas Kofoid and Campbell

Xystonellopsis cyclas Kofoid and Campbell, 1929, p. 244, fig.
457-

The fairly short lorica, of general cymatica type, with
thickened cylindrical pedicel and long lance, has a length of
5.0 oral diameters. The oral margin is thin and spread. The
bowl has a ringlike (0.25 oral diameter in width) ledge
(1.15 oral diameters in diameter) located just below the
anterior 0.1 oral diameter. The bowl below this ledge tapers
subconically (8° in the anterior 0.39 and 25 in the aboral
0.17 total length). The pedicel is subcylindrical, with a
diameter near 0.3 oral diameter and with a length of 0.32
total length. At its aboral end is a slightly flaring skirt.
From the center of the skirt arises the conical (15 ), sharply
pointed lance, the length of which is 0.14 total length.

The wall reaches a thickness of nearly 0.2 oral diameter at
the ledge, and is half as much just below; it thins down in
the lower bowl. There are very thin laminae and enclosed
radial secondary prisms in a single layer. In the ledge are
small, crowded alveoles in several rows. In the lance are
large hexagons in two layers. The surface has neat hex-
agonal structure, and there are 6 short pleats above the skirt.
The lumen does not enter the ledge, and in the lance it is
reduced to a canal.

Length, 238 to 249IX.

Xystonellopsis cyclas, like X. crassispinosa, has a thickened,
subcylindrical pedicel, but the lance is longer and more
slender, and the ledge more ringlike. It differs from X.
cymatica in having a thicker, more cylindrical pedicel and
thicker ledge.

Recorded from three stations in the Pacific, as follows:
one (156) in the Pacific equatorial region, and two (158,
160) in the region of South Pacific island fields.

There are 3 net samples, all taken at 100 meters. Fre-
quency, 4 per cent at stations 156, 160; average, 3.3 per cent.

Temperature, 26?62-28?52 (27^65); salinity, 35.06-35.89
(35-54); density, 22.76-23.14 (22.93); P H » 8 -34- 8 -44 ( 8 -3 Q )-

Xystonellopsis cymatica (Brandt) Jorgensen emended
Kofoid and Campbell
(Figure 65)
Xystonellopsis cymatica, Kofoid and Campbell, 1929, p. 245,

fig. 458.
The relatively short lorica, with wide upper ledge, bare

secondary ledge, conical bowl, thin pedicel, and short lance,
has a length of 4.82 oral diameters. The thin oral margin
is smooth. The bowl has a low upper cylindrical section
about 0.1 oral diameter in width, and the flat-edged, hori-
zontal ledge arises from its lower end. This upper ledge has
a diameter of 1.37 oral diameters, and its lower surface con-
tracts with concave contour to the bowl below. The lateral
wall within a length of 0.7 oral diameter is distinctly con-
cave. At the lower end arises a secondary ledge, with a
diameter of 1.25 oral diameters. Below this less distinct
ledge the bowl rapidly resumes its concave-conical form (20
in the upper 0.6 total length and 33 in the lower 0.4 total
length), with diameter at the aboral end, at the upper end
of the pedicel, only 0.16 oral diameter. The pedicel is narrow
and tubular, with a length of 0.37 total length, with some
slight skirtlike expansion at its distal end, and with little
lateral concavity in some individuals. From the lower end
of the skirt arises the narrow conical (7 ), pointed lance,
with a length of 0.58 oral diameter.

The wall is thickest under the upper ledge, where it
reaches nearly 0.13 oral diameter; in the lower bowl it is
about three-tenths as much. There are thin, hyaline laminae
with enclosed radial secondary prisms in a single row, save
suborally, where there are several layers of small hexagons.
The lumen does not enter the ledges, but does follow the
outer contour of the skirt, and in the lance forms a narrow
canal.

Length, 180 to 2581X.

A part of the Carnegie loricae conform exactly to the
typical form, but a good many, like the figured specimen
(fig. 65), differ in two respects: (a) in the presence of a
secondary ledge, and (b) in the presence of a slight skirt
below a thinner pedicel. These differences are not, in our
opinion, sufficient to justify designating a new species, at
least with the material at hand.

Xystonellopsis cymatica lacks the thick suboral angular
swelling of X. spicata, the more robust pedicel of X. cyclas
and X. crassispinosa, and the extremely narrow posterior
cone of X. mascarensis. In no case is the secondary ledge
like that of X. dicymatica.

Recorded from twelve stations, five in the Atlantic and
seven in the Pacific, as follows: one (19) in the Sargasso
Sea, four (22, 23, 25, 27) in the Atlantic equatorial region,
one (41) in the Galapagos region, five (86, 92, 93, 96, 98) in
the region of South Pacific island fields, and one (no) in
the North Pacific trade region.

There are 7 pump and 6 net samples, of which 2 were
taken at 50 meters and 11 at 100 meters. Frequency, 3
loricae in a pump sample at station no; average in pump
samples, 1.2 loricae.

Temperature: Atlantic, net samples i4?6o-22?42 (i8?68);
Pacific, pump samples i4?55~27?65 (23?85). Salinity:
Atlantic, net samples 35.70-37.05 (36.15); Pacific, pump
samples 34.73-36.22 (35.52). Density: Atlantic, net samples
25.30-26.62 (25.98); Pacific, pump samples 22.75-26.11
(23.55). pH: Atlantic, net samples 7.93-8.25 (8.09); Pacific,
pump samples 7.92-8.28 (8.17).

TAXONOMY AND DISTRIBUTION

83

Xystonellopsis dahli (Brandt) Kofoid and Campbell

Xystonellopsis dahli, Kofoid and Campbell, 1929, p. 245, fig.
483.

The rather elongated lorica, with little-differentiated aboral
region and without a skirt, has a length of 6.15 oral diam-
eters. The oral margin is thin and barely spread. The long
bowl is subcylindrical in the upper three-fifths, and sub-
conical (23 ) below. The pedicel is subcylindrical, 0.19 total
length in length, and without a skirt. The lance is short,
conical (2 ), 0.54 oral diameter in length, and sharply
pointed at the free tip.

The wall has a thickness of nearly 0.08 oral diameter; it
has thin laminae and enclosed fine hexagons; the outer sur-
face is almost clear, but there are 6 subvertical lists on the
pedicel.

Length, 396 to 444U.

Xystonellopsis dahli is shorter, with less aboral concavity
and less aboral differentiation than X. \rammeri. It has no
skirt as do X. tenuirostris and related species.

Recorded from one station (95) in the region of South
Pacific island fields, in 1 pump and 1 net sample, both taken
at 100 meters. Frequency, 2 per cent.

Temperature, 28^74; salinity, 35.35; density, 22.43; P H >
8.22.

Xystonellopsis dicymatica (Brandt) Kofoid and Campbell

Xystonellopsis dicymatica, Kofoid and Campbell, 1929, p. 245
(for fig. 467 see X. inaequalis).

The moderately tall lorica, with two wide, equal rings
marking the ends of a suboral spool, and with conical bowl,
thin pedicel, and narrow lance, has a length of 5.7 oral diam-
eters. The oral margin is thin and erect. The thickened
suboral spool occupies the anterior oral diameter. Just below
the oral rim is the upper ring, with a diameter of 1.4 oral
diameters; the lower one has a diameter not exceeding 1.5
oral diameters. The diameter of the bowl midway between
the rings is 1.3 oral diameters. The bowl below the spool
contracts conically (16 in the anterior 0.23 and 20° in the
posterior 0.28 total length). The pedicel is very thin (less
than 0.2 oral diameter at the most) and fairly short (0.29
total length), and has an expanded, aborally truncated skirt.
The lance is conical (2°), 0.6 oral diameter in length, and
sharply pointed.

The wall has a thickness of not over 0.1 oral diameter in
the lower bowl. There are thin laminae, which enclose
several layers of small hexagons in the spool and a single
layer of radial rectangles in the lower bowl; in the skirt
region there are several layers of tiny alveoles. The lumen
does not enter the rings and in the lance is reduced to a fine
canal.

Length, 250 to 280U.

Kofoid and Campbell (1929, fig. 467) have a lorica with
a wide lower ring nearly 1.6 oral diameters across. This
specimen is better assigned to Xystonellopsis inaequalis,
though the ring is not saucer-shaped as in the usual in-
aequalis.

Xystonellopsis dicymatica differs from X. inaequalis in
having the two rings subequal. U has 2 rings, whereas X.
cymatica has but one, anil A', abbreviata has three.

Recorded from nineteen stations, ten in the Atlantic and
nine in the Pacific, as follows: one (19) in the Sargasso Sea,
five (22, 23, 24, 27, 28) in the Atlantic equatorial region,
four (31, 32, 33, 34) in the Caribbean Sua, two (35, 154) in
the Pacific equatorial region, two (1.1, (17) in the South
Pacific middle latitudes, two (68, ~^) in the Galapagos
region, one (u$) in the North Pacific middle latitudes, and
two (147, 148) in the California region.

There are 1 pump and 21 net samples, oi which 5 were
taken at 50 meters and 17 at 100 meters. Maximum fre-
quency, 15 per cent at station 19; other records above mini-
mum (2 to 15 per cent) from stations 19, 28, 32, 33, 35, 61,
67, 148, 154; averages in net samples, 2.9 and 4.0 per cent in
the Atlantic and Pacific, respectively.

Temperature: Atlantic, net samples i7?34~26?79 (22^32);
Pacific, pump sample 21 "74, net samples io?92-25?8i
(i7?96). Salinity: Atlantic, net samples 36.00-37.05 (36.34);
Pacific, pump sample 34.66, net samples 34.05-35.47 (34.84).
Density: Atlantic, net samples 23.98-26.29 (25.15); Pacific,
pump sample 24.06, net samples 22.98-26.06 (25.11). pH:
Atlantic, net samples 7.99-8.30 (8.17); Pacific, pump sample
8.23, net samples 7.92-8.39 (8.07).

Xystonellopsis favata (Brandt) Jorgensen

Xystonellopsis favata, Kofoid and Campbell, 1929, p. 240, fig.
477-

The fairly elongated lorica, with wide bowl, short pedicel,
and lance, has a length of 4.0 oral diameters. The oral margin
is thin and erect. The bowl swells within the anterior 0.3
oral diameter to 1.08 oral diameters and then contracts (21 °)
for 0.5 oral diameter, forming, within the upper region, a
thickened section. Below this thick region the bowl is sub-
conical (8° in the anterior 1.25 oral diameters and 33 in
the lower 1.0). The pedicel is short (0.75 oral diameter),
tubular, and squarely truncated aborally. From its lower
end arises the conical (io°) lance, the length of which is
0.42 oral diameter; its distal tip is sharply pointed.

The wall averages approximately 0.08 oral diameter in
thickness in the bowl. There are thin laminae, enclosing
several layers of extremely faint, minute alveoles. The
lumen follows the contour of the bowl, but in the pedicel it
forms a narrow canal which continues to the tip of the lance.
On the pedicel are 12 subvertical lines or folds.

Length, 220 to 290(1.

Xystonellopsis favata resembles X. scyphium save that its
bowl is relatively longer and narrower, and its pedicel is less
set off from the bowl. It is not distant from X. tenuirostris,
but its bowl is relatively wider, with more aboral taper, and
the pedicel is not so stout or so long.

Recorded from one station (75) in the Galapagos region,
in a net sample taken at 100 meters. Frequency, 3 per cent.

Temperature, i8?40; salinity, 35.47; density, 25.55; pH,
8.10.

84

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Xystonellopsis gaussi (Laackmann) Kofoid and Campbell
Xystonellopsis gaussi, Kofoid and Campbell, 1929, pp. 246-247,

fig. 480.
The moderately elongated lorica, with armors-like form,
has a narrow lance and a length of 5.23 oral diameters. The
oral margin is thin and erect. The elongated bowl is sub-
conical (3° in the anterior 0.43 and 23 in the posterior 0.27
total length). The pedicel tapers (12°), has a length of 0.2s
total length, and terminates in a squarely truncated, non-
expanded skirt. The lance is pencil-shaped (15 ), and 0.5
oral diameter in length.

The wall reaches 0.07 oral diameter in thickness in the
bowl, and has thin laminae which enclose faint, minute
alveoles. There are 8 right-wound (3 ) discontinuous folds
on the pedicel.

Length, 371 to 47011.

Xystonellopsis gaussi has a pencil-like lance, rather than
a wide, conical lance as in X. armata. It differs from X,
acuminata in size, proportions, and presence of a distinct
pedicel; from both of these species it differs in having dis-
continuous folds rather than ridges on the pedicel. It has a
definite skirt, unlike X. heros and related species.

Recorded from eight stations, two in the Atlantic and six
in the Pacific, as follows: two (2, 16) in the Gulf Stream, two
(35, 153) in the Pacific equatorial region, one (63) in the
South Pacific middle latitudes, one (75) in the Galapagos
region, and two ( 109, 151) in the North Pacific trade region.
There are 4 pump and 9 net samples, of which 1 was
taken at the surface, 4 at 50 meters, and 8 at 100 meters.
Maximum frequency, 9 per cent at station 2; other records
above minimum (2 to 7 per cent) from stations 35, 63, 75,
151; averages, 4.2 per cent in Atlantic net samples, 4.0 per
cent and 3 loricae in Pacific net and pump samples, respec-
tively.

Temperature: Atlantic, net samples 20?50-23%4 (2i?26);
Pacific, pump samples i5?84-22?73 (i8?84), net samples
i 4 ?33-i 9 ?8i (i7?22). Salinity: Atlantic, net samples 36.40-
36.43 (36.41); Pacific, pump samples 34.42-34.76 (34.62),
net samples 34.86-35.47 (35.03). Density: Atlantic, net
samples 24.84-25.76 (25.50); Pacific, pump samples 23.83-
25.48 (24.76), net samples 24.72-26.00 (25.48). pH: At-
lantic, net samples 8.16-8.23 (8.20); Pacific, pump samples
8.08-8.31 (8.15), net samples 7.88-8.18 (8.02).

Xystonellopsis hastata (Biedermann) Kofoid and Campbell
(Figure 70)

Xystonellopsis hastata, Kofoid and Campbell, 1929, p. 247, fig.
456.

The fairly short lorica, with oral teeth, conical bowl, and
short pedicel and lance, has a length of 3.66 oral diameters.
The oral margin has 24 equidistant, short, triangular den-
ticles. The bowl expands slightly (1.13 oral diameters)
within 0.15 oral diameter, then contracts (3 in the anterior
1. 14 oral diameters and 34 in the posterior similar length);
the diameter at the lower end of the bowl is 0.3 oral diam-
eter, and the whole lorica is generally convex, with a suboral
ledge. The pedicel contracts from the lower end of the bowl

(io°), has a length of 1.0 oral diameter, and at its lower
end has a short, expanded skirt. From the center of the
skirt arises the short (0.4 oral diameter), fairly stout, pointed
lance.

The wall reaches 0.1 oral diameter in thickness across the
ledge where it is widest; it thins down to three-tenths as
much in the middle and lower bowl. There are thin laminae
with enclosed radial, secondary, rectangular prisms, with
minute alveoles within each one in several layers. The sur-
face has large hexagonal areas, except for the lance, which is
hyaline. There are 7 (8) short folds above the skirt on the
lower pedicel. The lower pedicel is yellowish, and denser
than the upper parts.
Length, 244H.

Xystonellopsis hastata is unique in the genus in having
oral denticles. In general it is related to X. cymatica and
relatives of that species, but is longer and stouter, with fuller
lower bowl, relatively shorter pedicel, and less localized sub-
oral thickening.

Recorded from five stations, three in the Atlantic and two
in the Pacific, as follows: three (23, 24, 25) in the Atlantic
equatorial region, and two (35, 152) in the Pacific equatorial
region.

There are 2 pump and 5 net samples, of which 1 was
taken at the surface and 6 at 100 meters. Maximum fre-
quency, 3 per cent at station 152; other records above mini-
mum (2 per cent) from stations 24, 35; average in Pacific
net samples, 1.2 per cent.

Temperature: Atlantic, pump samples 1 5?55-27? 1 8(21 ^36),
net samples i4?6o-i5?55 (i5?07); Pacific, net samples
n?48-i4?33 (i2?9o). Salinity: Atlantic, pump samples
35.22-35.61 (35.41), net samples 35.61-36.02 (35.83); Pacific,
net samples 34.73-34.91 (34.82). Density: Atlantic, pump
samples 22.84-26.34 (24.59), net samples 26.34-26.62 (26.44);
Pacific, net samples 26.06-26.50 (26.28). pH: Atlantic,
pump samples 7.96-8.32 (8.14), net samples 7.93-8.18 (7.99);
Pacific, net samples 7.76-7.88 (7.82).

Xystonellopsis heroica Kofoid and Campbell
Xystonellopsis heroica Kofoid and Campbell, 1929, p. 247, fig.

473-
The rather short lorica, of extreme slenderness and elon-
gation, has a length of 7.4 oral diameters. The oral margin
is thin. The bowl is swollen to a maximum of 1.17 oral
diameters within the upper oral diameter. Below this
swollen region it is subconical (4° in the anterior 0.37 and
30° in the posterior 0.21 total length). The pedicel-lance
has a length of almost 0.33 total length and is distally sharply
pointed.

The wall scarcely reaches 0.1 oral diameter in thickness in
the thickened region and is elsewhere less than a third as
much. There are thin laminae with enclosed fine hexagons.
There are no lists or ridges and the whole lorica is glass-
clear. The lumen enters the pedicel as a canal.
Length, 174 to 21811.

Xystonellopsis heroica has a bowl ami pedicel-lance of
greater elongation than in X. pinnata and X. heros. which

TAXONOMY AND DISTRIBUTION

85

species it otherwise rather closely resembles. Were it lack
ing in suboral thickening, it would be close to Parundella
longa.

Recorded from one station (94) in the region of South
Pacific island fields, in a pump sample taken at 100 meters.
Frequency, minimum.

Temperature, 28?66; salinity, 3547; density. 22.56; pH,
8.21.

Xystonellopsis heros (Cleve) Kofoid and Campbell

Xystonellopsis heros, Kofoid and Campbell, 1929, p. 247, fig.
484.

The very tall lorica, with distinct division between pedicel
and lance, has a length of 7.9 oral diameters. The oral rim
is flattened. The long bowl tapers (5 ) directly from the
nm without local modification throughout its whole length,
and reaches 0.3 oral diameter at the squarely truncated aboral
end, which is the homologue of the skirt of other species.
The lance is narrow conical (2 ), 0.58 oral diameter in
length, and sharp-pointed at the free tip.

The wall has a subuniform thickness of 0.08 oral diam-
eter, has thin laminae with enclosed faint hexagons, and the
surface is glass-clear save for a few irregular, discontinuous,
subvertical lines down the lower three-tenths. The lumen
enters the lance as a canal.
Length, 475^.

Xystonellopsis heros is elongated like X. lyammeri, but
has no skirt. Xystonellopsis acuminata has a plumper bowl
with distinct pedicel region, as do X. gaussi, X. lenuirostrts,
and X. armata.

Xystonellopsis heros, as here described, hardly resembles
the figure given by Brandt (1906), differing in length, ab-
sence of folds on the bowl, and thinner lance. It is probably
the same as the species designated by Kofoid and Campbell
(1939) as X. tropica, new species. If so, the distributional
data in this report include records of both species.

Recorded from twenty-one stations, twelve in the Atlantic
and nine in the Pacific, as follows: two ( 14, 16) in the Gulf
Stream, two (17, 18) in the Sargasso Sea, five (24, 25, 27,
28, 29) in the Atlantic equatorial region, three (31, 33, 34)
in the Caribbean Sea, one (75) in the Galapagos region,
three (109, 139, 151) in the North Pacific trade region, one
(113) in the North Pacific middle latitudes, two (136, 147)
in the California region, and two (152, 153) in the Pacific
equatorial region.

There are 7 pump and 21 net samples, of which 7 were
taken at 50 meters and 21 at 100 meters. Maximum fre-
quency, 34 per cent at station 151; other records above mini-
mum (2 to 19 per cent) from stations 14, 16, 27, 29, 31, 33,
34, 75, 136, 147, 152, 153; averages in net samples, 1.5 and
1 1.3 per cent in the Atlantic and Pacific, respectively.

Temperature: Atlantic, pumpsamples i8?o8-23?64 (22^44),
net samples i4?6o-26?04 (>o?63); Pacific, i9?8i-23?77
(22^02) and i4?32-22?73 (i8?64), respectively. Salinity:
Atlantic, pump samples 36.03-36.48 (36.30), net samples
35.59-36.82 (36.32); Pacific, 34.59-35.18 (34.82) and 34.42-
35.47 (34.86), respectively. Density: Atlantic, pump samples

24.84-26.06 (25.6$), net samples 23.98-26.66 (25.56); Pacific,
23.42-24.72 (24.08) and 23.83-25.75 (25.01), respectively.
pH: Atlantic, pump samples 8.09-8.23 (8.18), net samples
7.93-8.27 (8.16); Pacific, 8.18-8.28 (8.23) and 7.87-8.39
(8.18), respectively.

Xystonellopsis inaequalis Kofoid and Campbell

Xystonellopsis inaequalis Kofoid and Campbell, 1929, pp. 247-

248, fig. 466.
Xystonellopsis dicymatica, Kofoid and Campbell, 1929, rifj. 467

(for p. 245 see X. dicymatica).

The fairly short lorica, of general dicymatica type, with
the lower ledge much wider than the upper one, has a length
of 7.0 oral diameters. The thin oral margin is erect. The
bowl forms a short cylinder within the anterior 0.25 oral
diameter, at the lower end of which arises the Hat, ringlikc
upper ledge, with a diameter of 1.28 oral diameters. Below
this ledge the suboral band swells with a wide sigmoid curve
to the secondary or lower ledge, the diameter of which is
1.68 oral diameters. The suboral band occupies the anterior
1.08 oral diameters. The bowl below the band is subcorneal
(14 increasing to 30 in the lower oral diameter). The
pedicel tapers (2 ), with a length of 0.39 total length, and
at its distal end is an expanded skirt. The free edge of this
skirt has 6(8) clawlike projections. From the center of the
skirt arises the pencil-like lance (0.64 oral diameter in
length).

The wall is thickest in the suboral band, where it reaches
as much as 0.2 oral diameter; elsewhere it is quickly reduced
to three-tenths as much or even less. There are thin, hyaline
laminae with enclosed radial secondary prisms in a single
layer, save in the band, where there are several layers of
smaller hexagons. The lumen does not enter the ledges.
The whole lorica, save the skirted region, is translucent; the
latter is brownish.
Length, 246 to 350ft.

The Carnegie loricae are more slender (7.0 oral diam-
eters as against 5.75) than usual; they are also longer (up to
35011). Most of them have a concave secondary ledge, this
being like a dish or saucer rather than a flat ring.

Xystonellopsis inaequalis resembles X. dicymatica closely
except that the lower ledge is much wider and the suboral
band is generally wider. Intergrades do not seem to be
frequent.

Recorded from eight stations, four each in the Atlantic
and the Pacific, as follows: three (18, 19, 20) in the Sargasso
Sea, one (34) in the Caribbean Sea, two (35, 37) in the
Pacific equatorial region, one (78) in the Galapagos region,
and one (82) in the region of South Pacific island fields.

There are 3 pump and 5 net samples, of which 3 were
taken at 50 meters and 5 at 100 meters. Frequency, 3 per
cent at station 19; 2 per cent at station 35; average in Atlantic
net samples, 1.2 per cent.

Temperature: Atlantic, net samples 20?32-24?63 (22?48);
Pacific, net sample i6?3o, pump samples i9?82-24?34
(22?o6). Salinity: Atlantic, net samples 36.55-37.05 (36.78);
Pacific, net sample 34.88, pump samples 34.53-36.46 (35.76).

86

OCEANIC TINTINNOINA OF LAST CRUISE OF CARNEGIE

Density: Atlantic, net samples 24.65-26.67 (25.69); Pacific,
net sample 25.60, pump samples 24.47-25.11 (24.74). P^ :
Atlantic, net samples 8.19-8.25 (8.21); Pacific, net sample
7.92, pump samples 8.00-8.19 (8.1 1).

Xystonellopsis krammeri (Brandt) Kofoid and Campbell

Xystonellopsis \rdmmeri, Kofoid and Campbell, 1929, p. 248,
fig. 485.

The greatly elongated lorica, with heros-like form and
with spiral lines on the pedicel, has a length of 8.9 oral
diameters. The oral margin is sharp, with a bare suboral
flare. The very long bowl is subconical (7 as a whole, but
2° in the upper two-thirds and io° in the aboral third). The
pedicel is subcylindrical and about 0.6 oral diameter in
length, with a scant skirt at its lower end. The lance is
wide conical (20°), and 0.6 oral diameter in length.

The wall has a thickness of less than 0.07 oral diameter.
There are thin laminae with enclosed minute alveoles in the
upper bowl; in the lower bowl and pedicel these become
large prisms. The pedicel has 8 slightly (5 ) left-turned
ridges which arise at the skirt and extend the whole length
of the pedicel.

Length, 535 to 61 8(1.

Xystonellopsis \riimmeri is closest to X. dahli, but is
much longer and has a clearer-cut skirt. It bears some re-
semblance to X. heros, but lacks the distinct lance, and is
much longer; it differs similarly from X. acuminata.

Recorded from six stations, three each in the Atlantic and
the Pacific, as follows: two (28, 30) in the Atlantic equa-
torial region, one (34) in the Caribbean Sea, one (67) in
the South Pacific middle latitudes, one (145) in the North
Pacific middle latitudes, and one (146) in the California
region.

There are 6 net samples, of which 1 was taken at 50 meters
and 5 at 100 meters. Maximum frequency, 7 per cent at
station 67; other records above minimum (2 per cent) from
stations 28, 145, 146; average in the Pacific, 3.7 per cent.

Temperature: Atlantic, 2i?02-25?54 (23?3o); Pacific,
i6?36-2o?o7 (i8?53). Salinity: Atlantic, 36.40-36.65 (36.56);
Pacific, 34.32-34.70 (34.44). Density: Atlantic, 24.25-25.76
(25.03); Pacific, 24.24-25.45 (24.72). pH: Atlantic, 8.16—
8.28 (8.22); Pacific, 8.09-8.34 (8.23).

Xystonellopsis laticincta Kofoid and Campbell

Xystonellopsis laticincta Kofoid and Campbell, 1929, p. 248,
fig. 4 6 9-

The moderately long lorica, with thick suboral band,
conical bowl, long pedicel, and flaring skirt, has a length of
6.9 oral diameters. The oral rim is thin and erect. The
suboral band occupies the anterior 0.21 total length. It has
an upper and a lower ring with diameters of 1.45 oral diam-
eters, and the included region is thickened. The bowl below
the spool is subconical (io° in the anterior 0.19 and 40 in
the posterior 0.13 total length). The pedicel is wide at its
upper end (0.55 oral diameter) and tapers (4 ) to just above
the flaring skirt. The skirt is repeated. From its middle

arises the lance. The lance is conical (2 ), needle-like, and
0.56 oral diameter in length.

The wall has a thickness of 0.1 oral diameter below the
suboral spool; in other regions it is like that of Xystonellopsis
abbreviata. There are left-turned striae above the skirt.

Length, 304 to 343(1.

Xystonellopsis laticincta has a wider suboral band and
longer pedicel than X. abbreviata. It is narrower, with a
shorter, lower bowl, thinner pedicel, and more slender lance
than X. torta. The pedicel is shorter, the skirt less suddenly
differentiated, and the spool wider than in X. clevei.

Recorded from two stations (86, 87) in the region of South
Pacific island fields, in 2 pump samples taken at 100 meters.
Frequency, 2 per cent at station 86; average, 1.5 per cent.

Temperature, 24?07-25?u (24^59); salinity, 36.02-36.22
(36.12); density, 24.25-24.41 (24.33); pH, 8.23-8.25 (8.24).

Xystonellopsis ornata ( Brandt) Kofoid and Campbell

(Figure 69)

Xystonellopsis ornata, Kofoid and Campbell, 1929, p. 249, fig.
463.

The rather large lorica, with remarkable spool-like upper
differentiation, conical bowl, and lower dagger-like mass,
has a length of 3.8 oral diameters. The oral rim is erect,
thin, and sharp. The upper bowl has unique differentiation
(analogous with that of Stelidiella simplex and partially with
that of Epicranella bella). There is a narrow angular ledge
(1.16 oral diameters) within the anterior 0.05 oral diameter,
followed by a wider angular but narrowed one (1.45 oral
diameters) at 0.84 oral diameter below the rim. The spool-
like region thus formed has 12 vertical, sinuous, equidistant
ribs, which arise from the upper surface of the wide ledge
and run decurrently to just below the upper ledge: these ribs
subdivide the upper bowl into 12 curved, subequal facets.
The bowl contracts (19 ) subconically for about 0.42 total
length below the lower ledge and at the lower end has a
diameter of approximately 0.5 oral diameter; its anterior
diameter just below the wide ledge is 0.84 oral diameter.
The remaining 0.38 total length is an irregular, dagger-like
mass homologous with the pedicel and the lance of other
species of Xystonellopsis. This mass spreads out (20°) to
0.55 oral diameter near 0.8 oral diameter from the aboral
tip, and below that level is more or less regularly reduced
subconically (42°) to the sharply pointed aboral tip. The
aboral end is minutely open.

The wall reaches almost 0.28 oral diameter in thickness
across the lower ledge, but its average thickness is hardly
0.05. There are thin laminae and enclosed faint radial
secondary prisms. The wall thickens up in the aboral 0.38
total length and leaves a narrow canal to descend to the open
tip; the canal has a widened region at about 0.75 of its
length. The secondary alveoles of this posterior region are
in several layers and are hexagons. The lumen does not
enter the ledges, being generally conical. The surface has a
clear but faint hexagonal meshwork.

The animal is rather small and has 2 macronuclei.

Length, 262(1.

TAXONOMY AND DISTRIBUTION

87

The Carnegie loricae are longer and also less full aborally
than those from other sources.

Xystonellopsis ornata can scarcely be confused with any
other Xystonellopsis, for none of them has the upper spool-
like structure, nor the aboral dagger-shaped mass. This
upper differentiation is possibly correlated with the relation
of the lorica to the water, and the lower mass may act as a
counterpoise in directed locomotion.

Recorded from three stations in the Pacific, as follows:
one (35) in the Pacific equatorial region, and two (41, 74)
in the Galapagos region.

There are 3 net samples, of which 1 was taken at 50
meters and 2 at 100 meters. Frequency, 6 per cent at station
41; average, 3.3 per cent.

Temperature, i4?55-i6?55 (i5?79); salinity, 34.88-35.14
(35.01); density, 25.60-26.11 (25.82); pH, 7.89-7.92 (7.91).

Xystonellopsis paradoxa (Cleve) Jorgensen
(Figure 63)
Xystonellopsis paradoxa, Kofoid and Campbell, 1929, p. 249,
fig- 463-

The fairly short lorica, with duplicated knob and slender
lance, has a length of 4.0 oral diameters. The oral margin is
sharp-edged and erect. The bowl expands (40 ) within the
anterior 0.35 oral diameter to a diameter of 1.23 oral diam-
eters, at that level forming a ledge. Below, the bowl con-
tracts, and then swells to 1.12 oral diameters at 0.58 oral
diameter below the rim. The swollen region continues over
a band of approximately 0.5 oral diameter, and then con-
tracts subconically (n°) for a length equal to 1.62 oral
diameters, below which it rapidly contracts (6o c ) for a
short distance. The pedicel begins flaring (75°), and reaches
0.77 oral diameter at 0.71 total length from the rim. The
lower edge of this skirt is crenulate, and left-turning, short
folds extend upward from the crinkles. The secondary skirt
arises at 0.5 oral diameter below the primary one. Its width
is similar to its length, and, like the former one, its edge is
crenulated. The lance (0.73 oral diameter in length) is nar-
row conical (9°), arises from a spreading base, and is pointed
at its free tip.

The wall is thickest in the suboral swollen region, where
it reaches 0.16 oral diameter; elsewhere it is only a tenth as
much. There are thin laminae which enclose radial second-
ary prisms. The lumen is generally conical in the bowl,
bu