University of Southern California Dissertations and Theses

Ultrastructure Of Tests Of Some Recent Benthic Hyaline Foraminifera

(USC Thesis Other)

Ultrastructure Of Tests Of Some Recent Benthic Hyaline Foraminifera

PDF

Download a page range

Download transcript

Copy asset link

Request this asset

Transcript (if available)

Content I l
70-25,06*1
STAPLETON, Richard Pierce, 1935-
ULTRASTRUCTURE OF TESTS OF SOME RECENT
BENTHIC HYALINE FORAMINIFERA.
University of Southern California, Ph.D.,
1970
Paleontology
| University Microfilms, A XEROX Company, Ann Arbor, Michigan
THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED
ULTRASTRUCTURE OP TESTS OP SOME RECENT
BENTHIC HYALINE PORAMINIPERA
by
Richard Pierce Stapleton
A Dissertation Presented to the
FACULTY OP THE GRADUATE SCHOOL
UNIVERSITY OP SOUTHERN CALIFORNIA
In Partial Fulfillment of the
Requirements for the Degree
DOCTOR OP PHILOSOPHY
(Geological Sciences)
June 1970
UNIVERSITY OF SOUTHERN CALIFORNIA
T H E G R A D U A T E S C H O O L
U N IV E R S IT Y PA R K
L O S A N G E L E S . C A L IF O R N IA S 0 0 0 7
This dissertation, written by
RICHARD PIERCE STAPLETON
under the direction of h.l .S.... Dissertation C om
mittee, and approved by all its members, has
been presented to and accepted by The Gradu
ate School, in partial fulfillment of require
ments of the degree of
D O C T O R O F P H I L O S O P H Y
Dean
CONTENTS
Chapter Page
ABSTRACT ............................................ x
INTRODUCTION ........................................ 3
PREVIOUS INVESTIGATIONS .......................... A
METHODS OP INVESTIGATION .......................... 12
ACKNOWLEDGMENTS................................... 18
DESCRIPTION OF SPECIMENS .......................... 19
Bollvlna argentea Cushman .................... 19
Bollvlna splssa Cushman ...................... 71
Bollvlna acuminata Natland .................... 87
Bollvlna aenarlensls (Costa) .................. 90
Bollvlna Inter .luncta Cushman......... .... 93
Bollvlna pseudobeyr1chi Cushman ............. 96
Bollvlna semlnuda Cushman .................... 99
Cancrls maorlcus Finlay ...................... 10A
Lentlcullna cultrata (Montfort) ............. 120
Marglnullna obesa Cushman non Terquem .... 123
Islandlella callfornlca (Cushman and Hughes) . 129
Islandlella lomltensls (Galloway and Wissler) 132
Islandlella tortuosa (Cushman and Hughes) . . 136
Florllus baslsplnatus (Cushman and Moyer) . . 138
11
Chapter Page
Nonlonella mlocenlca Stella Cushman and Moyer 151
Cassldullna aff. C. laevigata d'Orblgny . . . 162
Cassldullna carlnata Silvestri ................... 167
Cassldullna pulchella d'Orblgny ................. 170
Cassldullna brazlllensls Cushman ................. 173
Pullenla bulloldes (d'Orblgny) ................... 178
Pullenla sallsburyl R. E. and K. C. Stewart . 181
Pullenla malklnae Coryell and Mossman .... 184
Chllostomella ovoldea Reuss . ................... 187
Qyroldlna neosoldanll Brotzen ................... 195
Hoeglundlna elegans (d'Orblgny) ................. 200
Stomatorblna concentrlca (Parker and Jones) . 212
ULTRASTRUCTURE OP BENTHIC HYALINE TESTS ............ 216
Comparisons between light and electron
microscopy....................................... 216
Grains and crystals.............................. 218
Organic matter ..................................... 219
P o r e s ............................................. 220
Laminae........................................... 221
The secondary calcite crust ............... . 222
Radial versus granular walls ..................... 225
The process of calcification..................... 226
ENVIRONMENTAL CONTROL OP ULTRASTRUCTURES ............ 229
ULTRASTRUCTURES AND GENETIC RELATIONSHIPS .... 231
ill
Chapter Page
SUMMARY AND CONCLUSIONS............................. 237
REFERENCES.......................................... 241
APPENDICES.......................................... 249
Appendix I: Sample localities .............. 250
Appendix II: Plate numbers ................... 255
iv
TEXT-FIGURE
Figure Page
1. Variations in the test morphology of
Bolivlna argentea Cushman .................. 21
v
LIST OP PLATES
Plate Page
1. Bollvlna spp. In cross-polarized light . . 23
2. Bollvlna argentea Cushman ............. 2^
3. Bollvlna argentea Cushman ............. 29
4. Bollvlna argentea Cushman ............. 32
5. Bollvlna argentea Cushman .................. 34
6. Bollvlna argentea Cushman .................. 36
7. Bollvlna argentea Cushman ..................
8. Bollvlna argentea Cushman .................. ^2
9. Bollvlna argentea Cushman .................. ^4
10. Bollvlna argentea Cushman .................. ^
11. Bollvlna argentea Cushman .................. 50
12. Bollvlna argentea Cushman .................. 52
13. Bollvlna argentea Cushman .................. 55
14. Bollvlna argentea Cushman ................ 57
15. Bollvlna argentea Cushman ................ 59
16. Bollvlna argentea Cushman ................ 61
17. Bollvlna argentea Cushman ................. 63
18. Bollvlna argentea Cushman . ............... ^5
19. Bollvlna splssa Cushman ................... 73
20. Bollvlna splssa Cushman ................... 75
21. Bollvlna splssa Cushman ................... 78
vl
Plate Page
22. Bollvlna splssa Cushman ............. . . . 80
23. Bollvlna splssa Cushman .................... 83
24. Bollvlna splssa Cushman .................... 85
25. Bollvlna acuminata Natland ................ 88
26. Bollvlna aenarlensls (Costa) .............. 91
27. Bollvlna Inter.lunota Cushman.............. 94
28. Bollvlna pseudobeyrlchl Cushman ........... 97
29. Bollvlna semlnuda Cushman .................. 100
30. Bollvlna semlnuda Cushman .................. 102
31. Foraminifera In cross-polarized light . . . 105
32. Cancrls maorlcus Finlay ..................... 108
33. Cancrls maorlcus Finlay ..................... Ill
34. Cancrls maorlcus Finlay ..................... 113
35. Cancrls maorlcus Finlay ................. . 116
36. Cancrls maorlcus Finlay ..................... 118
37. Lentlcullna cultrata (Montfort) ............ 121
38. Marglnullna obesa Cushman non Terquem . . . 125
39. Marglnullna obesa Cushman non Terquem . . . 127
40. Islandlella spp. in cross-polarized light . 130
41. Islandlella callfornlca (Cushman and
Hughes) . ................................. 133
42. Islandlella lomltensls (Calloway and
Wlssler) T ................................. 136
43. Foraminifera In cross-polarized light . . . 139
vll
Plate Page
44. Ielandlella tortuosa (Cushman and Hughes) . 141
45. Florllus baslsplnatus (Cushman and Moyer) . 144
46. Florllus baslsplnatus (Cushman and Moyer) . 147
47. Florllus baslsplnatus (Cushman and Moyer) . 149
48. Nonlonella mlocenlca Stella Cushman and
Moyer........................................ 152
49. Nonlonella mlocenlca Stella Cushman and
Moyer........................................ 155
50. Nonlonella mlocenlca stella Cushman and
Moyer........................................ 157
51. Nonlonella mlocenlca Stella Cushman and
Moyer........................................ 160
52. Nonlonella mlocenlca stella Cushman and
Moyer........................................ 163
53. Cassldullna spp. In cross-polarized light . 165
54. Cassldullna aff. C. laevigata d'Orblgny . . 168
55. Cassldullna carlnata Sllvestrl ........... 171
56. Cassldullna pulchella d'Orblgny ........... 174
57. Foraminifera In cross-polarized light . . . 176
58. Cassldullna brazlllensls Cushman ......... 179
59. Pullenla bulloldes d'Orblgny ............. 182
60. Pullenla sallsburyl R. E. and K. C. Stewart 185
61. Foraminifera in cross-polarized light . . . 188
62. Pullenla malklnae Coryell and Mossman . . . 190
63. Chllostomella ovoldea Reuss ............... 193
vlii
Plate Page
64. Foraminifera in cross-polarized light . . . 196
65* Gyroldina neosoldanii Brotzen .............. 198
66. Hoeglundina elegans (d'Orbigny) ............ 203
67. Hoeglundina elegans (d'Orbigny) ............ 205
68. Hoeglundina elegans (d'Orbigny) ............ 208
69. Hoeglundina elegans (d'Orbigny) ............ 210
70. Stomatorblna concentrica (Parker and Jones) 213
iz
ABSTRACT
Tests of twenty-six species from thirteen genera of
Holocene benthic hyaline foraminifera were examined by
means of light and electron microscopy. Six of these
species were examined In detail In order to determine the
extent to which ultrastructure Is controlled by environment.
The other twenty species were chosen In order to determine
the extent of genetic control. The ultrastructure of tests
Is controlled by both environmental and genetic factors.
A secondary calclte crust develops on tests of most
benthic hyaline species. With increasing depth, this crust
increases In thickness and, In doing so, produces a series
of characteristic surface configurations: (1) in shallow
water, a thin layer of calclte rhombs averaging 0.1 microns
across; (2) a pattern of zigzag grooves separating angular
areas about one micron across; (3) an irregular surface of
high relief; and (4) at the lower end of the depth range, a
pattern of sinuous grooves separating irregular areas
several microns across. The extent to which the crust
develops, and the depths at which the different surface
configurations occur, vary with different species. In
deep basins of low oxygen concentrations, development of
the crust Is Inhibited.
x
The degree to which the test wall Is laminated may
Increase, or decrease, or remain about the same with In
creasing depths, depending on the species. An Increase In
the degree of lamination results In smaller and more regu
lar grains.
Ultrastructure of tests Is consistent within species
when allowance Is made for changes produced by environmental
factors. Among some genera, such as Bollvlna, there was
little resemblance between test ultrastructure of con
generic species; congeneric species of other genera, such
as Oassldullna and Pullenla, had very similar ultrastruc
ture. Similarities were also noted In ultrastructure of
species from the related genera Nonlonella, Florllus.
Pullenla and Nonlonelllna. The ultrastructural features
which are most likely to be consistent within genera or
higher taxa are the size and shape of pores, and the appear
ance of walls In cross-sections.
On deep water species of Pullenla and Cassldullna.
distances between pores are two to three times as great as
on congeneric species from shallower water.
Test walls are constructed of monocrystalline grains
from one half to four microns In diameter. In radial walled
species, the (0001) planes of crystals have a preferred
orientation parallel to wall surfaces. In granular walled
species, the (1011) planes are parallel to surfaces. Cer-
xl
tain genera, such as Olblcldes and Gyroldlna. have a wall
type which is intermediate in its optical properties be
tween radial and granular. Optical properties of walls are
consistent within species and probably within most higher
taxa.
Under the light microscope, some species have a
fibrous appearance to test walls and many species have
nonfibrous outer layers. These phenomena are due to an
organic matrix which occurs as a three dimensional reti
culum between the grains. Pores are produced as a result
of the process of calcification.
Recrystalllzatlon can produce extreme changes in
the ultrastructure of fossil specimens within a relatively
short period of time. Hence, it is not likely that ultra
structure can be of great use for paleoecologlcal investi
gations or for systematic studies of fossil species.
xli
INTRODUCTION
I
Although relatively few papers have been published
on electron microscopic investigations of foraminiferal
tests, these studies Indicate that a considerable amount
|
of variation occurs in the appearances at high magnifica
tions of tests of different species. These variations in
test ultrastructure can be attributed to two possible
factors— genetics and environment, the latter including
i dlagenesis. The problem is to determine to what extent
| ultrastructure is controlled by each of these two factors.
This in turn suggests the further problems of whether or
not ultrastructure can be used for systematic purposes
i and whether or not it can be used for paleoecological
I
Investigations.
There are a number of subsidiary areas of investiga
tion in which ultrastructural studies may be of use. Al
though those optical phenomena which are observed when thin
i
; sections of tests are examined in polarized light have
been used for systematic purposes for some years now, the
nature of these phenomena is little understood. Crystal
sizes are too small to be adequately studied with light
microscopes and such studies are hampered by the thick
nesses of the sections which are usually many times thicker
than the diameters of the individual crystals which make up
tests. Investigators in many different fields are lnterest-
1 _
ed in the processes of calcification, yet few extensive
studies of organically precipitated calcium carbonate at
high magnifications have been published. Some Indications
of the size and arrangement of carbonate grains may be of
use to such Investigations.
The present study was restricted to Holocene species
of the benthic hyaline group. It must be recognized that
any attempt to carry out a thorough Investigation of test
ultrastructure would be a most formidable task. There are
over twenty thousand known species of foraminifera, of
which three quarters occur only as fossils, many poorly
preserved, and of which many are extremely rare. Some such
restriction is necessary, therefore, in order to obtain
results which are not dispersed over so wide an area as to
be of little value. Of the groups not studied, the plank-
tonic foraminifera are characterized by a number of micro-
structural features which, because of their high relief,
can be more adequately examined by scanning electron micro
scopy rather than by the transmission microscopy used in
this investigation. By eliminating fossil species, it was
possible to eliminate the factor of recrystallization
which, as will be shown, can produce radical and erratic
changes in ultrastructure within a relatively short period
of time. Previous investigations have shown that the ultra
structure of porcelaneous tests is quite different from
that of the hyaline forms and can best be studied in an
Independent investigation. Finally, the non-calcareous
foraminifera would require different investigative tech
niques and problems of their ultrastructures would be of a
different nature.
Twenty-six species, representing thirteen genera,
were chosen out of this selected group— the Holocene benthic
hyaline foraminifera— for investigation. Although to a
certain extent the choice of Bpecles was controlled by the
availability of a sufficient number of specimens, the
species used are reasonably representative of the entire
group. Six of the species were investigated in consider
able detail in order to study changes produced by varia
tions in the environment. The remaining twenty species
were chosen for information on the amount of control
exercised by genetic factors.
These twenty-six species represent less than 1 per
cent of all Holocene benthic hyaline species. Consequently,
even though the species were selected to be representative
of this group, it is necessary to use caution when extend
ing results to cover the entire group.
4
PREVIOUS INVESTIGATIONS
Interest In the microstructures of the calcareous
foraminifera dates back to Williamson (1858) who separated
the imperforate or "porcelainous" (^sicj foraminifera from
the perforate or hyaline forms. This separation was based
largely on examinations of thin sections under polarized
light, which demonstrated that there was a fundamental
structural difference between the two groups. It was not
baBed on the presence or absence of pores, a more super
ficial characteristic.
Sorby (1879) described tests of hyaline foraminifera
as consisting of minute prisms of calclte oriented with
the principle crystallographic axes perpendicular to the
surface of the test. When chambers are observed in cross
polarized light, each chamber shows a black cross sur
rounded by colored rings. Sorby also noted that some
species did not show a symmetrical arrangement of calclte
crystals but rather that the tests were constructed of
crystals arranged in small symmetrical groups. In a few
species, the wall of the test was constructed of granules
without any definite optical orientation.
A considerable number of species was examined under
cross polarized light by Wood (19^9) who divided the
hyaline foraminifera into two groups— those with test walls
constructed of radially arranged calclte crystals, and
5
those In which the crystals showed no preferred orientation
and thus produced a granular appearance to the test. Wood
considered this distinction to be of some systematic im
portance although he noted the apparent anomaly that
Ehrenberglna hystrlx Brady had a radial wall, while
Ehrenberglna hystrlx glabra Heron-Alien and Earland had a
granular wall.
In a later paper, Wood and Haynes (1957) noted that
Clblcldes refulgens Montfort had a radial wall although
originally reported by Wood (19^9) as being granular. The
authors did not give the source for this error but expressed
caution over use of the radial-granular dichotomy for
systematic purposes.
Krasheninnikov (1956) subdivided the two wall types
as follows:
Radial walls:
(a) Thin radial (e.g., Oassldullna sp.) with
walls composed of crystals 0.3 to 1.0 microns in diameter
oriented perpendicular to the surface;
(b) Coarse radial (e.g., Ammonia beecarl1
(Linne)) with walls composed of crystals 2.0 to 3.0 microns
in diameter;
(c) Indistinct radial (e.g., Elnhldlum sp.)
with walls composed of vermicular crystals which produce a
wavy extinction.
Granular walla:
(a) Jagged granular (e.g., Nonlon spp.) with
walls composed of equant grains 1.0 to 1.5 microns In
diameter;
(to) Lamellar granular (e.g., Olblcldes loba-
tulus (Walker and Jacob)) similar to the Jagged granular
wall but with three distinct layers and with larger grains;
(c) Mlcrogranular (e.g., CanalIfera sp.) with
crystals variable In size and shape.
It Is apparent that the genus Olblcldes Montfort
presents some peculiarities. Olblcldes refulgens was
reported as granular by Wood (1949) and C. lobatulus was
reported as granular by Krasheninnikov (1956). Wood and
Haynes (1957) then reported C. refulgens as radial while
the entire genus was described as radial by Loebllch and
Tappan (1964). Finally, Towe and Clfelli (1967) described
the walls of C. refulgens as indistinctly radial.
Krasheninnikov (1956) expressed the opinion that
the radial-granular division might be of use for systematic
purposes but was noncommltal on the use of subdivisions of
the two groups.
The genus Islandlella N/rfrvang was erected by N/tfr-
vang (1958) from species which had been previously placed
in the genus Cassldullna d'Orbigny. N^rvang separated the
two genera on the basis of apertural characteristics and
toothplates, and also noted that the walls of Islandlella
7
were radial while those of Cassldullna were granular. He
expressed caution on the use of this latter characteristic,
however.
Although these earlier authors approached with
caution the use of optical properties of the wall for
systematic purposes, in the most recent major classifica
tion of the foraminifera, that of Loeblich and Tappan
(1964), wall structure was used as a criterion for classi
fication as high as the subordinal level.
The type of wall in a specimen can be determined in
any one of three ways. The test may be sectioned and ex
tinctions observed directly; the entire test may be
examined for the pseudo-uniaxial cross figure; or the test
may be crushed in Immersion oil and the resulting frag
ments examined for the cross figure or for a preferred
extinction. This last is the simplest and quickest method.
Wood (1949) used both thin sections and crushed specimens
but does not state the method used for each species.
Krasheninnikov (1956) used thin sections, while N/frvang
(1958) does not state his methods. Loeblich and Tappan
(1964) used crushed specimens exclusively.
The first electron microscopic study of foraminifera
was that of Jahn (1953) who decalcified a number of cal
careous foraminifera and examined the resulting organic
material, particularly the organic linings of pores and
the sieve plates. The species examined were not identified.
8
Hay, et al. (1963) examined test surfaces by means
of direct carbon replicas and attempted to correlate their
observations with observations made at lower magnifications
with the light microscope.
Two papers, Hyde and Krinsley (1964) and Krinsley
and Be (1965), described techniques for electron microscopy
of foraminifera, although no examples were given.
The first attempt to put electron microscopy to a
✓
practical use in the study of foraminifera was that of Be
(1965) who used surface replicas to study the development
of a secondary calclte crust on a planktonic species
Globlgerlnoldes saccullfer (Brady). Be" concluded that
Bphaeroldlnella dehlscens (Parker and Jones) was a varia
tion of Globlgerlnoldes saccullfer in which a thick
secondary crust had produced an extreme modification of
the external morphology. This crust developed on indivi
duals living at greater depths.
A thorough study of the planktonic species Globoro-
✓
talla menardll (d'Orbigny) was carried out by Be, et. al.
(1966). The authors established the presence of a second
ary crust on this species also. This paper is particularly
noteworthy for the large number of excellent electron
micrographs which are figured.
Towe and Cifelli (1967) investigated a number of
benthic species by means of both surface replicas and
replicas of cross-sections. Although primarily concerned
9
with the process of calcification, a number of other topics
were discussed briefly, and the present author acknowledges
a considerable debt to this most informative paper. The
authors concluded that the granular wall was probably a
result of crystal development in which a (1011) plane was
oriented parallel to the surface of the wall, while in
radial walls, the crystals developed with the (0001) plane
paralleling the surface. Otherwise wall structures are
similar in the two groups and the division into radial and
granular wall types should not be used as a basis for major
taxonomic divisions. A model for the process of calcifica
tion was proposed and fundamental differences in the
structures of hyaline and porcelaneous walls were observed.
They also noted the presence of a thin veneer of calclte
overlying the outer surface of the test of Ammonia beccarll
✓
(Llnne) although they did not refer to this veneer as a
secondary calclte crust.
Lynts and Pfister (1967) studied surfaces of ten
benthic species. These authors also were concerned with
the process of calcification and with the organic linings
of pores. The presence of a thin calclte layer over sur
faces of tests of some porcelaneous species was noted.
Towe (1967) studied the cement and grain relation
ships in the walls of an arenaceous species Haplophragmoldes
canarlensls (d'Orbigny).
A technique for examining cross-sections of tests
10
was described by Hansen (1967).
Four papers, all published within the same year,
considered the possibilities of the scanning electron
microscope In micropaleontology: Hay and Sandberg (1967),
Hon Jo and Berggren (1967), Sandberg and Hay (1967), and
Bartlett (1967). The general Impression of all of these
authors was that the scanning electron microscope will be
of great benefit for studies of three dimensional structures
such as apertures and surface ornamentation, and for
studies of surfaces of planktonlc species In which the re
lief Is often too great for replication and transmission
microscopy.
Orr (1967), using light microscopy, studied changes
In the secondary calclte crust of four species of the
planktonlc genus Globorotalla Cushman. He found that the
crust developed In all four species below a certain depth,
each species having Its own peculiar depth at which crustal
development was Initiated.
Pessagno and Mlyano (1968) used electron microscopy,
phase contrast microscopy and dark field Illumination to
study wall structure of two planktonlc species, Globoro-
talia truncatullnoldes (d'Orbigny) and Globorotalla crassa-
formls (Galloway and Wissler). They described the septal
walls of these species as being granular rather than radial
but used these terms In a structural sense, not In a
crystallographlc sense, noting that the septa do show a
11
radial extinction In cross polarized light.
Towe and Cifelll (1967) had previously noted the
somewhat ambiguous use of the terms "radial" and "granular",
pointing out that ever since Wood's (19^9) study the terms
had generally been used to describe crystallographlc
orientation. They strongly recommended that the terms be
restricted to optical phenomena and not be construed to
have morphological or structural Implications. The
ambiguity of the terms has resulted In a number of authors
assuming that a radial wall must necessarily have a radial
structure while granular walls have a granular structure.
These authors noted that this Is not necessarily the case.
In the present paper, the terms will be restricted to
optical phenomena unless clearly stated otherwise.
The aragonitic species, Hoeglundina elegans
(d'Orblgny) was examined by Reiss and Schneldermann (1969)*
These authors were Interested In the process of calcifica
tion but Included a thorough description of the ultra
structures of this species. Since H. elegans was also
investigated in the present study, further reference will
be made to this paper below under the description of that
species.
12
METHODS OP INVESTIGATION
Many of the papers which have dealt with electron
microscopy of foraminiferal tests have included descrip
tions of the techniques which were used. The methods used
in the present study are variations on previously reported
techniques which in turn are developments of standard
methods of electron microscopy. There is a certain amount
of advantage to these present techniques in that they ap
pear to provide a large number of usable specimens within
a reasonable amount of time and with a minimum of effort.
Because of the low penetrating power of the electron
beam, specimens must be extremely thin, of the order of a
few hundred angstroms. Towe and Cifelli (1967) attempted
to section tests with an ultramicrotome but without suc
cess, probably because of the perfect cleavage of calclte
which makes cutting grains extremely difficult. These
authors consequently used replica methods as have all other
Investigators.
In replica methods, a thin film is prepared which
accurately reproduces the relief of a surface to be
examined. Such surfaces may be either natural, such as
the outer surface of a test, or artificial, such as the
plane produced by cutting through the test wall.
Test surfaces were examined by means of direct, or
single stage replicas. The tests are cleaned lightly by
15
brushing with a wet brush. More vigorous cleaning methods,
such as the use of organic solvents, strong oxidants, or
ultrasonic generators tend to alter the appearance of the
surface. The tests are placed on a glass slide with the
surface to be examined facing upward. In a vacuum eva
porator, a thin film of carbon is deposited perpendicular
to the slide on this surface. The carbon is followed by a
film of chromium evaporated at about a 30° angle to the
plane of the slide. The metal enhances relief because of
the shadowing effect produced by Irregularities on the
surface. Because of the curvature of the test, the angle
of shadow is not constant and consequently the breadth of
shadowed areas cannot be used to determine the amount of
relief. A second carbon layer is added over the chromium.
The resulting film is a sandwich of chromium between two
carbon layers and is not only extremely tough but is also
resistant to attack by the acid which is used to dissolve
the test.
Nickel specimen grids are placed on a glass slide
and a small amount (ca. 50 mg) of paraffin wax is placed
on each grid. The glass slide is heated until the wax
melts and coats the grids. Tests are placed, one on each
grid, with the carbon films facing down. The slide is re
heated until the wax melts a second time and when the slide
is cooled the tests are held to the grids by thin layers of
wax.
14
Tests are dissolved by placing the slide In 1.0
Normal hydrochloric acid. After about 20 minutes, any re
maining test material can be removed from the grids by
flicking lightly with a teasing needle. At this point,
only the carbon film remains, bonded to the grid by a thin
layer of wax.
The grids are gently lifted from the slide and
placed on small pieces of screening in a glass petri dish.
The dish is placed in an oven and left for three days at a
temperature of 100° C. During this time, the wax
evaporates allowing the films to settle onto the grids.
Finally, the grids are rinsed first in xylene to remove
any remaining wax, then in distilled water. They are
checked for quality in a light microscope and are ready for
viewing. Usually about 90 percent of the grids have good
films, with about 80 percent of each film usable.
In order to examine tests in cross-section, it was
necessary to use two stage, or double replicas. Each test
is embedded in a small block of epoxy resin. The block is
ground to produce a flat plane which cuts through the test
at the desired orientation. This plane is polished, then
etched for 15 seconds in 0.01 Molar EDTA. Primary
replicas are produced by pressing this surface against a
small square of heated polystyrene. The cross-section of
the test is accurately reproduced on the polystyrene.
These primary replicas are coated with a carbon and chromium
15
film similar to the films used for direct replicas of
surfaces.
A circle about 2 mm In diameter Is scored on the
coated polystyrene to enclose the area In which the test
Is replicated. The polystyrene is placed In a small dish
of xylene. After about 30 seconds, a teasing needle can
be carefully inserted at the edge between the film and the
polystyrene and used to free the scored circle. This
circle of carbon-chromium film is allowed to drift about
in the xylene for a few minutes while it is gently stroked
with a teasing needle to remove any remaining polystyrene.
It is then picked up on a copper specimen grid.
After the grids have dried, they are examined for
film quality in a light microscope and are ready for view
ing. Since four to five primary polystyrene replicas can
be made from each original specimen, there is usually at
least one good film for each original. Generally, the
second and third replicas are the best, since the first
replicas often are contaminated by extracted calcite or
organic matter, while later replicas are subject to the
formation of air spaces.
There was little apparent difference between
sections cut parallel to the long axis of the test and
those cut perpendicular to this axis.
Tests of four species were disaggregated in an
ultrasonic generator in order to examine the resulting
16
particles. Approximately twenty specimens of each species
were selected for lack of contamination by extraneous cal-
clte. These tests were placed In small vials of alcohol
which were then Immersed In an ultrasonic bath until the
tests disintegrated. Support films for the particles were
prepared by coating clean glass slides with an evaporated
carbon film. A few drops of the alcohol suspensions were
placed on the coated slides and allowed to dry. The
alcohol spreads as it dries, to give a fairly uniform dis
persion of particles across the coated slide. Chromium
was evaporated onto the slides at a low angle in order to
shadow the dispersed particles. The carbon film was then
scored into small squares which were floated off the slides
in distilled water where they were picked up on specimen
grids. When the grids had dried, they were ready for
viewing. Selected area diffraction was used on many
particles.
Thin sections of tests for examination in the
polarizing microscope were produced by Imbedding the tests
in blocks of epoxy resin and grinding the section from
these blocks. This method is considerably easier, faster
and more satisfactory than those in general use. As is
the case with replicas of cross-sections, the direction in
which the section was oriented was not of any great im
portance, although radial extinctions were somewhat
sharper in sections cut perpendicular to the long axis of
17
the test than those cut In other directions.
Specimens used In this Investigation were taken from
the collections In the Micropaleontology Laboratory of the
Department of Geological Sciences, University of Southern
California. Recent material was used In all cases, except
where otherwise stated, and whenever possible specimens
were used which had taken up the Rose Bengal stain, which
indicates that the animal was alive or only recently dead
when collected. Between four and eight specimens were
used for surface replicas and six to eight specimens were
used for cross-section replicas for each sample of each
species. The plates illustrate representative areaB on
these specimens. In most cases, there was little variation
among tests of the same species from each sample.
The fine structures were studied on an RCA EMU3F
Instrument in the Electron Microscopy Laboratory of the
Department of Biological Sciences and Allan Hancock
Foundation, University of Southern California.
Most of the original negatives were taken at a
magnification of 8000. On replicas of cross-sections, the
discontinuity between the test wall and the embedding
plastic often produces a fold or break in the replica.
This disrupted area, along with regions which replicate the
embedding plastic, has been blacked out on the plates.
18
ACKNOWLEDGMENTS
The author wishes to express his sincere thanks to
Dr. 0. L. Bandy, University of Southern California, who
suggested and encouraged this Investigation, and to Dr.
R. P. Blls, University of Southern California, who greatly
assisted the author In his work with the electron micro
scope. The author also acknowledges debts to Dr. M. J.
Evans and Mr. W. Mayr of the Electron Microscopy Laboratory
of the Department of Biological Sciences and Allan Hancock
Foundation where much of the work was carried out. The
remainder of the Investigation was undertaken In the Micro-
paleontology Laboratory, Department of Geological Sciences
and Allan Hancock Foundation, University of Southern Cali
fornia, which supplied working space, samples, equipment,
and much helpful discussion from the author's colleagues.
The manuscript was critically read by Dr. 0. L. Bandy, Dr.
R. F. Blls, and Dr. W. H. Easton, University of Southern
California. Financial assistance was provided by grants
GA-730 and GB-6628 from the U. S. National Science Founda
tion.
19
DESCRIPTIONS OP SPECIMENS
Bollvlna argentea Cushman
Bollvlna argentea Cushman is one of the more common
and abundant species off the coast of southern California.
The optimum depth range for the species 1b from 350 m to
1200 m, although specimens occur both above and below this
range. Within this depth range, the species is often the
dominant foraminiferal component of bottom samples.
The gross morphology of the test shows a consider
able amount of variation and this variation was investi
gated statistically by Lutze (1962, 1964). He found that
test morphology of this species ranged from a triangular,
keeled, costate form, often with a basal spine, to a more
rectangular form lacking basal spine or keel, and with
costae poorly developed or absent. The range of morphology
is so great that individual specimens have been placed in
as many as three different species and two subspecies.
Nevertheless, Lutze convincingly demonstrated that the
group is in fact a single species.
In shallow water, specimens are generally triangular,
keeled, costate forms and the extremely rectangular, un
ornamented forms are absent. With increasing depth, there
is a gradual shift to rectangular shapes until at the lower
end of the depth range, most specimens are from the rec
20
tangular end of the morphological range. Samples taken by
Lutze at intermediate depths from environments depleted In
oxygen showed an abrupt shift to the rectangular shape.
The variation in test morphology with depth and with oxygen
concentration is shown dlagrammatically in text-figure 1.
Wall thickness is from 5 to 15/u. In thin sections,
very thin dark lines are seen paralleling the surface and
are visible in both normal and cross-polarized light.
Under cross-polarized light, walls become extinct when
rotated into positions parallel to the directions of
polarization (pi. 1, figs. 1-2). Tangential sections under
cross-polarized light show dark bands of extinction across
chambers (pi. 1, fig. 3). The pseudo-uniaxial cross figure
is not usually seen in its entirety but the dark bands ap
pear to be arms of the cross figure. There were no dif
ferences among thin sections of specimens taken from dif
ferent depths or of different morphologies. Extinction in
cross-sections is the same for walls of early and late
chambers, and for septa, except on the proloculus where
the divisions between extinct and nonextinct areas is
abrupt, giving a grainy appearance, although the extinc
tion remains radial.
Because of the great morphological range in this
species, two groups of specimens were examined from each
sample, chosen to represent the extremes of variation in
that sample. For convenience, these groups will be desig-
TEXT-FIGURE 1
Variations In the test morphology of Bollvlna
argentea Cushman with depth and with oxygen
concentrations. Figures are adapted from
Lutze (1964).
21
TEST
MORPHOLOGY
LOW OXYGEN
PLATE 1
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-2 Bollvlna argentea Cushman
Thin section in two orientations at
45° to show the radial extinction.
3 Bollvlna argentea Cushman
Tangential section. Dark bands on
the chambers, indicated by arrows,
are arms of the pseudo-uniaxial
cross figure.
4-5 Bollvlna splssa Cushman
Thin section in two orientations at
45° to show the radial extinction.
23

25
nated as triangular from a deep sample.
Since the final chamber Is sometimes only partially
developed on some specimens, and hence may not be repre
sentative, the penultimate or antepenultimate chambers are
used to show characteristics of late chambers. For early
chambers, one of the first three or four pairs was used.
Plate 2 shows the ultrastructure of the surfaces of
a late chamber and an early chamber of a specimen of
triangular form from a depth of 357 m. On the late chamber
(pi. 2, fig. 1), pores 3.5 to 4.0/u in extreme diameter may
be seen. From this extreme diameter, the surface slopes
gently Inward until the diameter has decreased to 2.0 /a.
At this point, the pore reaches its minimum diameter, a
diameter which remains constant throughout the remainder of
the wall. This initial funnel-shaped entrance to the pore
is about 1 /U deep and will be referred to as the "pore
funnel."
Distances between pores vary from 6 to 10 /a. The
surfaces between the pores are made up of very small
rhombic faces which average about 0.05/u across. These
enclosed areas suggest euhedral calcite crystals. The
texture of the interpore regions extends down the sides of
the pore funnels and across the pore openings, Indicating
a calcite layer whioh coats the surface of the test and
covers the pores themselves. Dark cloudy regions In the
pore openings are organic matter and openings Into the
PLATE 2
1-2 Bollvlna argentea Cushman
Electron micrographs. Triangular form
from 357 m depth. 1, surface of a late
chamber; P, pore; PF, pore funnel; Z,
zigzag groove pattern. 2, surface of
an early chamber. X 20,000.
26
• T2&
f e ■ :
27
1
28
pores may exist, obscured by this material.
On the early chamber (pi. 2, fig. 2), no pores are
visible. The surface is extremely smooth and is almost
featureless. Since pores occur on early chambers of this
species, the thin calcite layer which was observed on the
late chamber must completely obscure the pores on early
chambers.
In plate 3* surfaces of late and early chambers of
rectangular forms from this same sample, from a depth of
357 m, are shown. On the late chamber (pi. 3» fig. 1)»
pore funnels are almost absent. Diameters of pores are
about 1.5/u and the pores appear to be open. Dark cloudy
material in the pore openings is organic matter. The dis
tances between pores are about the same as for the tri
angular forms although the area of the Interpore regions
is greater because of the absence of pore funnels. Inter
pore regions are similar in appearance to those of the
triangular forms except that the rhombic faces are either
absent or too small to be clearly shown, while the pattern
of zigzag grooves occurs over all of the Interpore regions.
On the early chamber (pi. 3» fig. 2), open pores can be
seen. The zigzag grooves are also clearly visible though
not as distinct as on the late chamber.
In cross-seotlon, walls of both triangular and
rectangular forms from this sample are similar, nor were
there any differences between walls of early and late
PLATE 3
1-2 Bollvlna argentea Cushman
Rectangular form from 357 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
29
30
31
chambers. Two cross-sections are shown In plate 4. Plate
4, figure 1, Is a cross-section of the wall of an early
chamber of a triangular form. The wall Is composed of Ir
regular, equant grains from 0.5 to 1.0 /a In diameter. It
Is not possible to determine crystallographlc orientation
on any of these grains. The structure indicated on the
figure Is a pore which Is filled with grains of calcite
similar In appearance to the grains In the remainder of
the wall. In general, there Is no pattern to the arrange
ment of the grains, though In some specimens a vague
laminar structure may be seen paralleling the surface of
the wall.
Plate 4, figure 2, Is a cross-section through a
costa on an early chamber of a rectangular form. Many of
the grains In this section have a rhombic shape suggesting
calcite faces with a common orientation. Toward the
Inner surface of the wall a vague laminar structure Is
visible.
In both of these figures, the Irregular granular
structure of the wall extends out to the outer surface
which Is generally smooth. Irregularities of the outer
surface which show up In these figures are largely arti
facts produced by the difficulties of obtaining good
replication across the discontinuity between the test wall
and the embedding plastic.
Plates 5 and 6 are surfaces of specimens from a
PLATE 4
1-2 Bollvlna argentea Cushman
Specimens from 357 m depth, 1, cross-
section of test wall; the outer surface
of the test Is toward the top of the
figure; P, pore; X 20,000. 2, cross-
section of test wall through a costa;
the outer surface of the test Is toward
the right of the figure; X 10,000.
32

PLATE 5
1-2 Bollvlna argentea Cushman
Triangular form from 463 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
34

PLATE 6
1-2 Bollvlna argentea Cushman
Rectangular form from 463 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
36
2
38
depth of 463 m. Late and early chambers of a triangular
form from this sample are shown In plate 5. The late
chamber (pi. 5» fig* 1) Is very similar In appearance to
the late chamber of the triangular specimen from 357 m.
However, the zigzag groove pattern Is not visible on the
specimen from the deeper sample and the rhombic faces are
considerably larger--up to 0.1/u in diameter. On this
figure, orientation of the rhombic faces can be seen to
vary across the surface of the chamber. The changes in
orientation are gradual and there are no lines of demarca
tion between regions of different orientation.
No pores are visible on the early chamber (pi. 5»
fig. 2). The surface is flat, covered with rhombic faces
averaging about 0.05 /u across. As on the late chamber,
orientation of the rhombs is not constant, although they
appear to be more distinct.
On the late chamber of a rectangular form from this
sample (pi. 6, fig. 1), pore funnels can be seen around the
pores but these pore funnels are smaller than those on the
triangular forms. The pores and pore funnels are almost
completely covered with a layer of calcite rhombs. These
rhombic faces appear to have a similar orientation. No
pores are visible on the early chamber (pi. 6, fig. 2) and
rhombic faces are considerably smaller than on late
chambers.
Specimens from a depth of 576 m are shown in plates
39
7 and 8. One pore, and possibly a second, can be seen on
the late chamber of a triangular specimen (pi. 7, fig. 1).
These pores are reduced to thin irregular slits but are
still open. Zigzag groove patterns on the surface separate
irregular, angular regions 0.2 to 0.6 /a across. To a con
siderable extent, this groove pattern resembles those seen
on specimens from 357 m depth. On the early chamber of a
triangular specimen (pi. 7, fig. 2), similar markings may
be seen though they are not so well defined.
Pores may be seen on the late chamber of a rect
angular specimen from 576 m depth (pi. 8, fig. 1) as slit
like openings. The surface is covered with the same type
of zigzag grooves as were observed on specimens from 357 m
and, as on these shallower specimens, the surface suggests
a mass of euhedral calclte crystals, each about 0.5/u in
diameter. The early chamber of the same specimen (pi. 8,
fig. 2) is very smooth with some features suggesting
indistinct rhombic faces from 0.1 to 0.2yu across.
The specimens shown in the next two plates, plates
9 and 10, are from a depth of 532 m in Santa Barbara Basin.
This sample was taken below the sill depth in the basin
and is from a region strongly depleted in oxygen concen
trations. Because of the shift in test morphology with
decreased oxygen, triangular forms from this sample cor
respond roughly in morphology with rectangular forms taken
from samples of the same depth but in regions of normal
PLATE 7
1-2 Bollvlna argentea Cushman
Triangular form from 576 m depth
1, surface of a late chamber; P,
2, surface of an early chamber.
X 20,000.
pore.
40
41
2
PLATE 8
1-2 Bollvlna argentea Cushman
Rectangular form from 576 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
42

PLATE 9
1-2 Bollvlna argentea Cushman
Triangular form from 532 m depth in a
low oxygen environment. 1, surface of
a late chamber. 2, surface of an early
chamber. X 20,000.
44

PLATE 10
1-2 Bollvlna argentea Cushman
Rectangular form from 532 m depth in
a low oxygen environment. 1, surface
of a late chamber. 2, surface of an
early chamber. X 20,000.
46
47
2
48
oxygen concentrations.
On late chambers of triangular forms from a depth
of 532 m in Santa Barbara Basin (pi. 9, fig* 1), pores
appear as subcircular regions filled with amorphous organic
matter. Poorly developed pore funnels may be seen around
some pores. Pores are about 3/u in diameter and are
spaced 6 to 10/u apart. The regions between pores are
extremely smooth with a faint speckled appearance which
suggests very small rhombic faces, too small for accurate
observation. In contrast, the interpore region of an
early chamber (pi. 9* fig* 2) is marked by sinuous grooves
deeply incised into the surface and enclosing irregular
areas from 0.5 to 1 . 5 across. The surfaces within these
enclosed areas are similar in appearance to the interpore
regions of the late chamber. Pores are visible on the
early chamber as irregular slits about 2/u long by 1
wide.
Late chambers of rectangular specimens from 532 m
depth (pi. 10, fig. 1) are similar in appearance to those
of the triangular specimens. Pores are smaller, about 2 yu
in diameter, and well developed pore funnels are visible.
Early chambers of this specimen (pi. 10, fig. 2) are al
most the same in appearance as late chambers except that
pore funnels are up to 3 fx in diameter and are exception
ally well developed. On both early and late chambers, the
Interpore regions are smooth and faintly speckled.
49
At a depth of 787 m, pores are no longer visible on
triangular forms. Plate 11, figure 1, shows a late chamber
of a triangular form from this depth. The surface has a
platy nature, clearly shown in this figure. An irregular,
angular area, relatively smooth, is surrounded by a higher
region of greater relief. Within this lower plate, rhombic
faces from 0.1 to 0.2 /i across are seen to maintain an al
most constant orientation. The remainder of the surface
is marked by coarse grooves delineating smooth areas which
vary in size up to 0.5 A1*
The surface of an early chamber (pi. 11, fig. 2)
consists of large flat areas 4 to 7 A1 across, separated by
irregular, deeply Incised grooves. The shadows indicated
on the figure show that these areas are of different eleva
tions.
Pores are still visible on rectangular forms from
787 m depth. On a late chamber (pi. 12, fig. 1), pore
funnels 2 to 3 A1 in diameter surround pore openings about
1.5A1 *n diameter. The interpore regions are dominated by
deeply incised grooves enclosing angular areas 0.5 to 1.0/\x
across. This surface coats the pore funnels and covers the
pore openings. An early chamber of one of these specimens
(pi. 12, fig. 2) has a very smooth surface although oc
casional grooves can be seen. Pores are not usually
visible and when present, are almost completely obscured
by the surface crust.
PLATE 11
1-2 Bollvlna argentea Cushman
Triangular form from 787 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber; note shadows
indicated by arrows. X 20,000.
50
51
2
PLATE 12
1-2 Bollvlna argentea Cushman
Rectangular form from 787 m depth
1, surface of a late chamber. 2,
face of an early chamber; Pf pore
X 20,000.
sur-
52

5*
Pores are rarely visible on either triangular or
rectangular forms from a depth of 842 m. The surfaces of
late chambers of triangular forms from this depth (pi. 13*
fig. 1) consist of smooth areas 0.5 to 2.5/u across
separated by zigzag grooves. Early chambers of the same
specimen (pi. 13* fig. 2) sure similar except that the
grooves are not as distinct. Late and early chambers of
rectangular forms from this sample (pi. 14, figs. 1-2) are
moderately smooth and are almost featureless although there
Is a slight suggestion of rhombic faces less than 0.1yu In
diameter.
The next four plates, plates 15 to 18, show surfaces
and cross-sections of specimens from 1050 m depth, near the
lower end of the depth range for this species. The surface
of a late chamber of a triangular form (pi. 15* fig. 1) Is
smooth and featureless except for a few sinuous grooves.
Early chambers of the same specimen (pi. 15* fig. 2) have a
higher relief and rhombic configurations can be seen over
most of the surface. Rhombic faces are about 0.1yu across
and maintain constant orientations within larger regions.
These larger regions, of very variable size, are separated
by shallow grooves. Surfaces of late chambers of rect
angular forms (pi. 16, fig. 1) are characterized by numerous
zigzag grooves enclosing smooth areas 0.3 to 0.5 yu across.
Early chambers of the same specimen (pi. 16, fig. 2) re
semble those of triangular specimens from this sample except
PLATE 13
1-2 Bollvlna argentea Cushman
Triangular form from 842 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
55
56
2
PLATE 14
1-2 Bollvlna argentea Cushman
Rectangular form from 842 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
57
58
2
PLATE 15
1-2 Bollvlna argentea Cushman
Triangular form from 1050 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
59
60
2
PLATE 16
1-2 Bollvlna argentea Cushman
Rectangular form from 1050 m depth.
1, surface of a late chamber. 2, sur
face of an early chamber. X 20,000.
61
2
PLATE 17
1-2 Bollvlna argentea Cushman
Triangular form from 1050 m depth.
1, cross section of wall of a late
chamber; the outer surface of the wall
Is toward the top of the figure; C,
secondary caicite crust. 2, cross-
section of wall of an early chamber;
the outer surface of the wall Is toward
the right of the figure; C, secondary
calclte crust; L, laminated region.
X 20,000.
63
64
2
PLATE 18
1-2 Bollvlna argentea Cushman
Rectangular form from 1050 m depth.
1, cross-section of wall of a late
chamber; the outer surface of the wall
Is toward the right of the figure; C,
secondary calclte crust; L, laminated
region. 2, cross-section of wall of
an early chamber; the outer surface of
the wall Is toward the right of the
figure; Ct secondary calclte crus; L,
laminated region. X 20,000.
65
66
67
that relief Is lower.
Plate 17• figure 1, Is a cross-section of the wall
of a late chamber of a triangular form. The outer surface
Is smooth and an outer layer 0.5/u thick may be seen. Be
low this outer layer, the test is constructed of irregular
grains about 0.5/U in diameter. Organic matter, seen as
cloudy dark masses, occurs between grains and between the
outer layer and the Inner part of the wall. There is a
vague laminar structure visible in the inner parts of the
wall.
In a cross-section of an early chamber of a tri
angular form (pi. 17, fig. 2), the innermost part of the
wall is formed of irregular equant grains 0.5 to 1.0 yu in
diameter. Between this region and the outermost layer,
there is a series of thin laminae totalling 3 to 5 /u in
thickness, each lamina being less than 0.4 u thick. These
laminae are each constructed of a single layer of grains,
each grain extending across the full thickness of the
lamina. The grains vary in length from 0.5 to 1.0/u and
are separated within the laminae by irregular boundaries.
Some organic matter can be seen between grains and between
laminae. Between this laminated region and the outer sur
face of the wall there is a layer slightly over 1 yu in
thickness in which individual grains are difficult to dis
tinguish.
The cross-section of the wall of a late chamber of
68
a rectangular form (pi. 18, fig. 1) Is similar In general
appearance to the wall of an early chamber of the tri
angular form. The laminated region is only about 2 yu thick
while the outer layer has a more definitely granular
structure. An early chamber of one of the rectangular
forms (pi. 18, fig. 2) Is again very similar. On none of
these cross-sections was it possible to determine the
crystallographlc orientation of the grains.
The total thickness of the test walls varies very
little between specimens from shallow water and those from
deep water. The laminated region of the test Is not,
therefore, a series of secondary layers but is rather a
different type of test construction to that used In shallow
water. A very poorly defined lamination Is seen on most
specimens and this lamination apparently becomes more
clearly defined with depth. A similar phenomenon will be
shown on other species which are unrelated to B. argentea.
The outermost layer, about 1 yu thick, which is found
on specimens from deep water is a secondary calclte crust,
analogous to the crust reported on so many planktonic
species. Generally, though not always, the crust is
thicker on early than on late chambers. This, along with
its progressive development from shallow to deep water,
indicates its secondary origin. Its occurrence on speci
mens which have taken up the Rose Bengal stain shows that
it is not a post mortem addition.
69
The thickness of the crust appears to Increase
gradually with Increases In depth. It does not form
abruptly at a certain specific depth as Orr (1967) noted
for planktonic species. Development of the crust from
shallow to deep water produces a series of characteristic
configurations on the surfaces of chambers. These con
figurations are best seen on late chambers as there is a
blurring of relief on early chambers, perhaps due to
abrasion or solution. At its Inception, near the upper end
of the depth range for the species, the crust appears as a
thin layer of calclte rhombs (pis. 5-6). These rhombs
show no preferred orientation over the surface of the
chamber, but may have a roughly constant orientation within
restricted areas. Pores are visible, though often partial
ly blocked by the crust, particularly on early chambers.
The next stage is the development of a zigzag groove pat
tern which delineates smooth, angular regions about 0.5/1
across (pi. 7» fig. 1; pi. 6, fig. 1). The grooves appear
to be grain boundaries separating single euhedral crystals
of calclte which have developed from the smaller rhombs.
At depths between 700 m and 900 m, surfaces develop a very
high relief and lack much of a definite pattern (pi. 11,
fig. 1). Below 900 m, the surface again becomes smooth and
is now separated into larger regions by sinuous grooves
(pi. 13, fig. 1; pi. 15* fig* 1). The sinuous grooves are
developments of the zigzag grooves but little can be ob
70
served of the crystallography of the enclosed areas.
The series of configurations does not follow a
perfectly regular pattern and there are numerous exceptions.
However, the general development holds true and these same
configurations will be seen on other species In which a
crust has developed.
The sample from a low oxygen environment (pis. 9-10)
shows very little crustal development, though In general
oxygen concentrations decrease with depth over the depth
range of this species. However, the oxygen concentrations
at 1050 m, the depth of the deepest specimens examined, are
not as low as those in Santa Barbara Basin below the sill
depth. In this same low oxygen environment, according to
Lutze (1964), there is a shift in test morphology toward
the rectangular forms, a shift similar to that which occurs
with increasing depth. However, while this shift is ac
companied by an increase in crust when it is produced by
increasing depth, the reverse applies when the shift is
produced by lowering of oxygen concentrations. It seems
likely, then, that depletion of oxygen operates independent
ly of depth in its control of crustal development.
Within each sample there is a very general tendency
for the crust to be less well developed on rectangular
forms than on triangular forms. In many caseB, pores on
rectangular forms are smaller and pore funnels are less
well developed.
71
Although thin sections of tests show a clearly
defined radial extinction under cross-polarized light,
there Is no corresponding radial structure to the wall.
Walls are constructed of Irregular, equant grains, the
crystallographlc orientation of which could not be deter
mined. Costae and keels are similar In structure to the
remainder of the wall and septa do not differ greatly
either. The laminated regions found In walls of some speci
mens from deep water do not noticeably affect the appear
ance of the test In cross-polarized light.
Bollvlna splssa Cushman
Bollvlna splssa Cushman Is another Holocene species
from off the coast of southern California. Its optimum
depth range Is somewhat greater than that of B. argentea—
from 300 to 1400 m. Lutze (1962) noted morphological
changes with depth In this species also, similar changes
to those which occur In B. argentea. Shallow water speci
mens are more triangular In shape, often keeled, with
costate early chambers, while deep water specimens are
slender, lacking keel or costae. The variations In
morphology, however, are neither as extreme nor as notice
able as those of B. argentea. Consequently, only those
specimens representing the median morphological form In
each sample were selected for examination.
Walls of this species are from 8 to 10yu In thick-
72
ness. In thin section* walls have an extinction similar to
that of B. argentea (pi. 1, figs. 4-5) although the dark
lines which were seen to parallel the surface of B.
argentea were not visible on B. splssa.
Surfaces of late and early chambers of a specimen
from 335 m depth are shown in plate 19. On the late
chamber (pi. 19* fig. 1)* the surface is formed of small
raised bosses* shaped like truncated cones* each about one-
third of a micron in diameter. Pores are from 2.5 to 3.0ja
in diameter, lacking the large pore funnels which were seen
on B. argentea. Distances between pores are from 6 to 10 ja»
The pores are open* but blocked by organic matter.
On the early chamber (pi. 19, fig. 2), the interpore
regions are similar except that the bosses are not as dis
tinct in this figure. Pores on early chambers are con
siderably larger, from 4 to 5yu in diameter. Very slight
pore funnels can be seen surrounding the pores* increasing
diameters by another micron. Distances between pores are
about the same.
In cross-section (pi. 20* fig. 1), the wall of a
late chamber of a specimen from this depth is formed of
three poorly defined but distinct laminae* the outermost
being the thinnest. A slight amount of organic matter
occurs between the laminae. The calclte in the wall has
very small rhombic faces* each about 0 . 0 5 across* which
have a constant orientation. It is not possible to detect
PLATE 19
1-2 Bollvlna splssa Cushman
Specimen from 335 m depth,
of a late chamber; P. pore,
face of an early chamber; P
X 20,000.
1, surface
2, sur-
pore.
73
74
PLATE 20
1-2 Bollvlna splssa Cushman
Speoimens from 335 m depth. 1, cross-
section of wall of a late chamber;
outer surface of the wall Is toward the
top of the figure. 2, cross section of
test wall at the Junction with a septum;
the outer surface of the wall Is toward
the right of the figure. X 20,000.
75

77
larger grains such as were seen In the walls of B.
argentea. A second cross-section was taken through the
junction of a septum with the wall (pi. 20y fig. 2). In
this region, the wall construction becomes more Irregular
and the laminae become obscured. Orientation of calclte
rhombs is constant over restricted areas but is no longer
constant throughout the wall.
Prom this depth, 335 m, to a depth of 576 m, there
is very little change in the ultrastructure. The surface
of a late chamber of a specimen from 576 m is shown in
plate 21, figure 1. The only noticeable change is a lower
ing of relief which causes the bosses to be less well de
fined.
At a depth of 787 m, the bosses have become still
less distinct, appearing as rounded features on the sur
face (pi. 21, fig. 2). Pores at this depth have developed
incipient pore funnels which are of the same diameter as
the entire pores are on shallower specimens. Openings into
pores have been reduced to 2 to 3yu in diameter.
At a depth of 988 m, the bosses are no longer
visible and calclte can be seen encroaching over one of the
pores (pi. 22, fig. 1). The surface relief is consider
ably lower. By 1050 m depth, the surface has become very
smooth and some pores are completely blocked by the
secondary calclte crust (pi. 22, fig. 2).
Prom 335 ® to 1050 m, the diameters of pores, in-
PLATE 21
1-2 Bollvlna splssa Cushman
1, surface of a late chamber of a speci
men from 576 m depth. 2, surface of a
late chamber of a specimen from 787 m
depth. X 20,000.
78
79
2
PLATE 22
1-2 Bollvlna splssa Cushman
1, surface of a late chamber of a speci
men from 988 m depth. 2, surface of a
late chamber of a specimen from 1050 m
depth. X 20,000.
80
81
2
82
eluding pore funnels, remained constant, with pores on
early chambers always larger than those on late chambers.
Pore spacing does not vary with depth.
At a depth of 1240 m, near the lower end of the
depth range for this species, zigzag grooves appear on the
surfaces of late chambers (pi. 23). On one specimen from
this depth (pi. 23i fig. 1)» a calclte layer with this
groove pattern can be seen closing a pore opening. On many
specimens, however, pores appear to be open, though blocked
by organic matter (pi. 23, fig. 2).
Cross-sections of walls of late chambers of speci
mens from this depth show no great changes from those of
walls of late chambers of specimens from shallow water (pi.
24). Although calclte rhombs are not as apparent on these
figures, this may be a result of the techniques used.
Orientation of the rhombic faces does not appear to be as
constant as on the shallow water specimens and there is an
indistinct radial structure.
The layer Indicated on the outer surface of the wall
in plate 24, figure 1, is different in structure from the
remainder of the wall and this layer is the same secondary
calclte crust as was seen on B. argentea. However, the
stage of development of the crust, with its zigzag groove
pattern and with pores still visible, occurs on B. argentea
in the upper third of its depth range, while on B. splssa
it does not develop until near the lower end of the depth
PLATE 23
Bollvlna splssa Cushman
Surfaces of late chambers of specimens
from 1240 m depth. X 20,000.
2
PLATE 24
1-2 Bollvlna splssa Cushman
Cross-sections of walls of late chambers
of specimens from 1240 m depth; the
arrow Indicates the secondary calclte
crust; outer surfaces of the walls are
toward the right of the figures.
X 20,000.
85

87
range for that species.
Bollvlna acuminata Natland
Specimens of Bollvlna acuminata Natland were col
lected off the coast of southern California at a depth of
12 m. This species Is generally confined to depths of
less than 200 m. Walls are 8 to 10 u thick and, In thin
section under cross-polarized light, resemble those of B.
splssa. Surface and cross-section views of late chambers
are shown in plate 25.
The pores are extremely large, over 5/U In diameter,
and contain a great deal of organic matter which makes ob
servation of pore characteristics difficult (pi. 25, fig.
1). Interpore regions have a relatively high relief with
calcite rhombs visible In many areas, the rhombs averaging
less than 0.2 yu across.
In cross-section (pi. 25, fig. 2), a series of thin
bands of calcite, separated by layers of organic matter,
are oriented almost perpendicular to the surface. Within
each of the calcite bands, crystallographic orientation
appears to remain constant.
It was not possible to determine whether or not
this species had a secondary calcite crust since many of
the features of the test were obscured by organic matter.
However, there does appear to be a surface layer about
1.5/u thick of different construction to the remainder of
PLATE 25
1-2 Bollvlna acuminata Natland
1, surface of a late chamber; the dark
region on the left Is a pore. 2, cross-
section of the wall of a late chamber;
the outer surface of the wall Is toward
the right of the figure. X 20,000.
88
89
2
90
the trail.
Bollvlna aenarlensle (Costa)
Bollvlna aenarlensls (Costa) Is very close to being
a sibling species of B. argentea. Geographic and depth
distributions of the two species are quite different and
B. aenarlensls does not seem to have the great amount of
morphological variation which Is found In B. argentea.
Specimens of B. aenarlensls are generally smaller and have
a greater curvature along the long axis. However, many
specimens of B. argentea would be extremely difficult to
distinguish from specimens of B. aenarlensls.
Test walls are about 10 yu thick. In thin section
under cross-polarized light, walls are similar in appear
ance to those of B. splssa.
The specimens used came from the Bay of Biscay from
a depth of 830 m. Late chambers are shown in plate 26 In
surface and cross-section views.
Pores are about 1.5 yu in diameter and pore funnels
are absent (pi. 26, fig. 1). Distances between pores vary
from 8 to 10/u• Zigzag grooves are visible over much of
the interpore regions. Overlying one of the pores in plate
26, figure 1, is a cloudy mass which Is an organic pore
lining. About half of the pores are open, the remainder
are covered with a secondary calcite crust.
In cross-section (pi. 26, fig. 2), the wall is seen
PLATE 26
1-2 Bollvlna aenarlensls (Costa)
1, surface of a late chamber; note pore
lining in lower left; P, pore. 2, cross-
section of the wall of a late chamber;
the outer surface of the wall Is toward
the right of the figure. X 20,000.
91

93
to be constructed of curved, platy grains, 0.5 to 1.0 yu
long by 0.1 to 0.2 yu thick, lying parallel to the surface.
1 secondary calcite crust of different texture and about
2 u thick may be seen in the figure. There is organic
matter between the grains. A number of indistinct laminae,
3 to 4 thick, can be seen in the wall.
Bollvlna inter.luncta Cushman
The specimens of Bollvlna inter.luncta Cushman were
collected off the west coast of South America at a depth of
420 m. This species is widespread throughout the eastern
portion of the Pacific Ocean. Walls are about 10^ thick
and, in thin section under cross-polarized light, resemble
walls of B. spls8a. Arms of the pseudo-uniaxial cross
figure are not usually seen in tengentlal sections because
of the very irregular surface of this species.
Surface and cross-section views of late chambers
are shown in plate 27. Pores are very small for this
genus, about 1 ^u in diameter, and are often blocked or
partially blocked by the secondary calcite crust (pi. 27,
fig. 1). Pores are spaced 5 to 7/i apart. In the inter
pore regions, the surface is composed of smooth flat areas
separated by deeply incised zigzag grooves.
The wall in cross-section (pi. 27, fig. 2) is com
posed of large irregular grains 1.0 to 1.5 across. Grain
boundaries are not well defined and all grains appear to
PLATE 27
1-2 Bollvlna Inter.luncta Cushman
1, surface of a late chamber; P, pore.
2, cross-section of the wall of a late
chamber; outer surface of the wall Is
toward the right of the figure.
X 20,000.
94

96
have a similar crystallographic orientation. No distinct
crust was visible In cross-section although a crust was
clearly visible in the surface view. A poorly defined
outer layer, 1 thick, may be the secondary calcite crust.
Bollvlna pseudobeyrlchl Cushman
Bollvlna pseudobeyrlchl Cushman is one of the most
delicate members of the genus, with walls under 4/u in
total thickness. It is found along the west coasts of the
American continents at depths from about 600 to 1000 m.
The specimens which were used came from a depth of 492 m
off the coast of southern California. In thin section
under cross-polarized light, the walls resemble those of
B. splssa although extinctions are difficult to observe
because of the very thin walls.
The surface of a late chamber is shown in plate 28,
figure 1. Pores are 2 to 3 yu in diameter and have very
slightly developed pore funnels. Spaclngs between pores
| are between 9 and 11 yU. Interpore regions are divided
into large smooth areas by deep, sinuous groove patterns
and faint rhombic faces are visible over most of these
areas. Most pores are open though partially blocked by
the secondary calcite crust.
In cross-section (pi. 28, fig. 2), the wall is com
posed of three to four well defined laminae, from 0.7 to
1.5/u thick. Each lamina is constructed of tabular grains
PLATE 28
1-2 Bollvlna pseudobeyrlchl Cushman
1, surface of a late chamber; P, pore.
2, cross-section of a late chamber; the
outer surface of the wall Is toward the
top of the figure, inner surface toward
the bottom. X 20,000.
97

99
which are bounded on two sides by the surfaces of the
lamina, but are divided from each other by irregular
boundaries within the lamina.
Bollvlna semlnuda Cushman
Bollvlna semlnuda Cushman is distributed throughout
the Pacific Ocean in depths from 150 to 1000 m. The speci
mens used came from the Gulf of California at a depth of
73? m. Test walls are relatively thin for the genus,
about 7 thick. In thin section under cross-polarized
light, walls resemble those of B. splssa.
The upper portions of chambers of this species are
free of pores and plate 29 shows both porous and pore-free
portions of a late chamber. Pores are large, 4.5 to 5.0
/U in diameter. Although many pores are open, over half
are either partially or completely closed by secondary
calcite crust. The interpore regions are composed of
flat, Irregular areas bounded by lightly Incised sinuous
grooves.
On the pore-free region (pi. 29, fig. 2), the
sinuous grooves are much more deeply Incised and the areas
bounded are generally larger than the areas in porous
regions. These pore-free regions resemble chambers of
B. argentea on which a very thick crust has developed (of.
pi. 15, fig. 1).
In cross-section (pi. 30), the wall is composed of
PLATE 29
1-2 Bollvlna semlnuda Cushman
1, surface of a porous area on a late
chamber; P, pore. 2, surface of a
pore-free area on a late chamber.
X 20,000.
100
101
2
PLATE 30
Bollvlna semlnuda Cushman
Cross-section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure; C, secondary calcite
crust. X 20,000.
102

104
a number of laminae of very variable thickness, from 0.5
to 3.0/u. Each lamina is formed of blocky grains which
extend across the full thickness of the lamina. Numerous
rhombic faces can be seen on these grains and, except in
the vicinity of pores, the grains appear to have a common
crystallographic orientation. The outermost lamina is the
secondary calcite crust which can be seen to have a dif
ferent type of construction.
Oancrls maorlcus Pinlay
The specimens of Cancrls maorlcus Pinlay which were
used in this study came from off the Pacific coast of
South America, from depths of 210 to 735 m. Although speci
mens are known to occur at greater depths, the numbers of
individuals in samples taken below 800 m decrease markedly.
Since large numbers of individuals were found in the
samples used, it would appear that these samples represent
the optimum depth range for this species.
Although a moderately large species, the walls of
C. maorlcus are only about 15 /i thick. In thin section
under cross-polarized light, a very strong extinction is
seen when walls are parallel to the directions of polari
zation (pi. 31» figs. 1-2). There is a somewhat fibrous
appearance to the walls in both normal and cross-polarized
light. Tangential sections and fragments of crushed tests
show a clearly defined pseudo-uniaxial cross figure (pi.
PLATE 31
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-3 Oancrls maorlcus Pinlay
1, thin section with wall at 45° to
the polarizers. 2, the same section
rotated until the wall Is parallel
to the polarizers so that the wall
becomes almost completely extinct.
3» tangential section showing a
broad pseudo-uniaxial cross figure.
4-6 Lentlcullna cultrata (Montfort)
4, thin section with wall at 45° to
the polarizers. 5» the same section
rotated until the wall Is parallel
to the polarizers so that the wall
becomes almost completely extinct.
6, a fragment of a test; extinction
produces a broad dark band across
the fragment.
105

107
31, fig. 3).
Most of the species examined In this investigation
showed similar surface configurations. The surfaces of
0. maorlcus, however, showed a considerable variation on
each chamber of each specimen and these configurations were
peculiar to this species. Near the keel, there are numerous
closely spaced pores. The numbers and sizes of pores de
crease from the keel down the side of the chamber toward
the umbilicus until the surface becomes pore free on the
large apertural flap.
In samples taken from depths of 210, 300 and 420 m,
there did not appear to be any great changes in surface
configurations between specimens taken from different
depths. However, the variations on each specimen made
comparisons of specimens from different depths difficult.
Plate 32, figure 1, shows a region on the surface
near the keel of the last chamber of a specimen from a
depth of 300 m. Pores are large open areas 2.5 to 3 , 0 /U
in diameter, spaced 4 to 6 pi apart. The narrow interpore
regions are formed of large irregular calcite crystals 1
to 2 jol in diameter, with similar orientations.
An area further away from the keel is shown in
plate 32, figure 2. This specimen came from a depth of
210 m. In this region of the test, pores are smaller,
about 1.5/a in diameter, and are spaced 8 to 10 p\ apart.
The interpore regions are formed of groups of radially
PLATE 32
1-2 Cancrla maorlcus Finlay
1, surface of a late chamber of a speci
men from 300 m depth showing a region of
large closely spaced pores. 2, surface
of a late chamber of a specimen from 210
m depth showing a region of smaller
pores, more widely spaced; F, fan
shaped group of crystals. X 20,000.
108

110
arranged crystals, each crystal about 0.1 by 0.3/a. In size.
There are flat fan-shaped groups of crystals extending
along one side of many of the pores. One such group Is
Indicated on the figure.
Still farther from the keel, these fan-shaped groups
of crystals extend across the pore to partially block the
pore opening (pi. 33* fig* 1)* At the base of the chamber,
near the umbilicus, pores are almost absent (pi. 33* fig.
2). Here, most of the surface appears to be formed of
Irregular or Indistinct rhombic faces averaging 0.2 /u
across. This configuration extends across the apertural
flap. Both figures In plate 33 are late chambers of
specimens from 210 m depth.
Cross-sections of late chambers of specimens from
a depth of 210 m are shown In plate 34. A region midway
between keel and umbilicus, free of large pores, Is shown
In plate 34, figure 1. The test wall Is constructed of
irregular grains 1 to 2 /a In diameter. Crystal faces on
many of the grains show a common crystallographic orienta
tion between grains. There is no apparent order to the
arrangement of the grains and no sign of the fibrous
structure which was seen In thin sections.
A region of large closely spaced pores Is shown In
cross-section In plate 34, figure 2. The pores extend
only about 5 /i into the wall where they are abruptly
truncated. Other "pores" begin and end within the test
PLATE 33
1-2 Cancrls maorlcus Finlay
1, surface of a late chamber of a speci
men from 210 m depth showing a region of
small pores. 2, surface of a late
chamber of a specimen from 210 m depth;
a pore-free region. X 20,000.
Ill

PLATE 34
1-2 Cancrls maorlcue Finlay
1, cross-section of the wall of a late
chamber of a specimen from 210 m depth
in a region of small pores; the outer
surface of the wall is toward the
right of the figure. 2, cross-section
of the wall of a late chamber of a
specimen from 210 m depth; 0, outer
surface to the right on the figure; P,
a large pore penetrating the wall part
way; IF, a second pore beginning and
ending within the wall. X 20,000.
113
2
115
wall with no apparent connection to either the outer or
the Inner surface of the wall. These "pores" produce a
sponge-llke texture to the wall In this region.
Specimens from a depth of 735 m show significant
changes from those taken In shallower water. Although
there was no noticeable change In regions of large pores,
a calclte crust Is visible on regions of small pores and
on pore-free regions. Plate 35» figure 1, Is the surface
of a final chamber In a region of small pores. The pore
openings are completely blocked by a secondary calclte
crust. Organic pore linings are visible on the surface
beside most pores. Relief in the Interpore regions Is
moderate and many rhombic faces with a common orientation
are visible. In a pore-free region (pi. 35* fig. 2), in
distinct sinuous grooves outline flat plates 2 to 3 ^
across.
In a cross-section of a pore-free region of a late
chamber on a specimen from this depth, the crust appears
as a distinct layer about 1.5 yu thick of different con
struction to the remainder of the wall (pi. 36* fig. 1).
Below the crust, the wall is similar to walls of specimens
from shallower water. On a cross-section of a region of
large pores (pi. 36* fig. 2), a distinct lamination of the
wall is visible. These laminae are not found on all cross-
sections but are fairly common in specimens from this
depth.
PLATE 35
1-2 Cancrls maorlous Finlay
1, surface of a late chamber of a speci
men from 735 m depth; a region of medium
pores; the dark mass extending from the
pore is an organic pore lining. 2,
pore-free region on the surface of a
late chamber of a specimen from 735 m
depth. X 20,000.
116
2
117
PLATE 36
1-2 Qancrls maorlcus Pinlay
1, cross-section of the wall of a late
chamber of a specimen from 735 «n depth
In a pore-free region; outer surface
of the wall Is toward the right of the
figure; C, secondary calclte crust. 2,
cross-section of the wall of a late
chamber of a specimen from 735 m depth
In a region of large, closely spaced
pores; 0, outer surface; P( pore.
X 20,000.
118

Lentlcullna cultrata (Montfort)
120
Lentlcullna cultrata (Montfort) is one of the
largest species of benthic hyaline foraminifera. Many
specimens attain a diameter of several millimeters with
walls up to 50/u thick. The specimens which were used
came from the southeastern Pacific at a depth of 866 m.
In thin section under cross-polarized light, there
is a pronounced radial extinction on walls which are
parallel to the directions of polarization (pi. 31 » figs.
4-5). Walls have a fibrous appearance, the fibers
oriented perpendicular to the wall surface, and an in
distinct laminar structure. At the outermost edge of the
wall is a thin banded layer about 8 yu thick which lacks
this fibrous structure. Tangential sections show a faint
preferred extinction which is better seen in fragments of
crushed tests. A fragment of a crushed test is shown in
plate 31, figure 6. A single broad dark band crosses the
center of the fragment parallel to one of the directions
of polarization. To the sides of this band, the fragment
is much lighter.
The surface of the test is smooth, marked by
closely spaced grooves which have a pattern midway be
tween zigzag and sinuous (pi. 37» fig* 1)* Areas enclosed
by the grooves are about 0.25/u across. Pores are widely
spaced and appear as irregular openings slightly under
0.5 /U in diameter, frequently filled with organic matter.
PLATE 37
1-2 Lentlcullna cultrata (Montfort)
1, surface of a late chamber; P, pore.
2, cross-section of the wall of a late
chamber; outer surface of the wall Is
toward the right of the figure.
X 20,000.
121

123
On some specimens, the grooves have a more sinuous pattern
and the enclosed areas are larger. On many of the speci
mens, scratches could be seen on the surfaces, possibly
produced by abrasion during processing of the sample.
In cross-section, walls are composed of Irregular
grains 1 to 3/u In diameter (pi. 37, fig. 2). There Is no
regularity to the arrangement of the grains and masses of
organic matter can be seen disseminated among the grains.
The calclte crust appears as an Indistinct outer layer.
In spite of the very fibrous appearance to the
test In thin section, and the strongly defined radial ex
tinction, there is no corresponding radial structure. Al
though the figure shows only the outer 5 /U of the wall,
the structure is similar throughout the wall from the
outer to the inner surface. The outermost layer which
could be seen In thin sections was not apparent under the
electron microscope.
Marelnullna obesa Cushman non Terquem
Like Lentloullna cultrata, Marglnullna obesa Cush
man non Terquem is an exceptionally large species. The two
genera are closely related members of the same family.
The sample from which specimens were taken was collected
in the Antarctic at a depth of 3800 m.
Test walls of this species are from 50 to 60 /i in
thickness. Under cross-polarized light, thin sections show
124
a very pronounced radial extinction (pi. 38, figs. 1-2).
The wall appears to be formed of a thick central layer of
fibrous structure bounded by two thinner layers of in
determinate structure. Tangential sections, in spite of
the great thickness of the wall, show a very clearly
defined pseudo-uniaxial cross figure (pi. 38, figs. 3-4).
Note that the cross becomes distorted in some orientations.
Surfaces of final chambers are marked by deeply
incised, extremely sinuous groove patterns (pi. 39* fig.
1). Areas bounded by the grooves are from 1 to 2^u across.
These areas are in turn divided into smaller areas by a
less distinct groove pattern. Pores are slightly over
0.25 yu in diameter, usually open and containing organic
matter. Distances between pores are from 2 to 3 /i.
In cross-section (pi. 39* fig. 2), the wall is
formed of a series of laminae which vary in thickness from
4 to 6 yu. Each lamina is constructed of a single layer of
blocky grains, each of which extends across the thickness
of the lamina, and which are 1 to 2 yu wide. Structure is
similar throughout the thickness of the wall. Although
grains terminate at the boundaries of the laminae, in
most cases grains in adjacent laminae are stacked over each
other. Organic matter occurs between grains and between
laminae.
Test construction is less regular in the outermost
few microns where grains are smaller and less blocky in
PLATE 38
1-4 Marglnullna obesa Cushman non Terquem
Cross-polarized light with polarizers
oriented parallel to the borders of the
plate. 1-2, thin section in two
orientations at 45° to show the radial
extinction; X 250. 3-4, tangential
section in two different orientations
to show the pseudo-uniaxial cross
figure; X 100.
125
126
/ *
£
0 -1 mm. i *
I 0 2 . m m I
! |
*
i 01 mm. i
PLATE 39
1-2 Marginullna obesa Cushman non Terquem
1, surface of a late chamber. 2,
cross-section of the wall of a late
chamber; outer surface of the wall is
toward the right of the figure.
X 20,000.
127
128
2
129
shape. In this outermost portion of the wall, larger
grains appear to be composed of smaller units with less
distinct boundaries. Since the larger grains are about
the same size as the areas on the surface bounded by the
well-defined grooves, while smaller units are about the
same size as the areas on the surface bounded by the less
well-defined grooves, It appears that the groove patterns
on test surfaces represent grain boundaries. A similar
correlation between grain sizes as observed In cross-
sections and groove patterns as seen on surfaces can be
observed In other species but Is seldom so clearly shown
as on specimens of M. obesa.
Islandlella callfornlca (Cushman and Hughes)
Specimens of Islandlella callfornlca (Cushman and
Hughes) were collected In 735 m depth of water off the
west coast of South America. Walls of this species
average about 30/a in thickness.
There is a very strong radial extinction observed
in thin sections under cross-polarized light (pi. 40, figs.
1-2). The wall has a thick fibrous central layer bounded
by two thinner layers of granular appearance but with
radial extinctions. In tangential section (pi. 40, fig.
3), the wall appears granular although broad diffuse
pseudo-uniaxial cross figures may be seen on some chambers.
Pores are visible on surfaces of late chambers as
PLATE 40
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-3 Islandlella callfornlca (Cushman and
Hughes)
1, thin section with wall at 45° to
the polarizers. 2, the same section
rotated until the wall is parallel to
the polarizers so that the wall be
comes extinct. 3* tangential section;
note the extinct areas on the chamber
to the upper left, appearing as two
arms of the pseudo-uniaxial cross
figure.
4-6 Islandlella lomltensls (Galloway and
Wissler)
1, thin section with wall at 45° to
the polarizers. 2, the same section
rotated until the wall is parallel to
the polarizers so that the wall be
comes extinct. 3» fragment of a
crushed test showing the pseudo-
uniaxial cross figure.
130
3
i 01 mm. i
6
132
small Irregular openings 0.5/u across separated by dis
tances of 3 to 6 /a (pi. 41, fig. 1). An examination of
several specimens from shallower water Indicated that
pores are normally about 1.5/u In diameter and that a
secondary calcite crust has partially blocked the pores on
these deeper specimens. Interpore regions are smooth,
composed of flat rhombic faces averaging about 0.25/U
across.
In cross-section (pi. 41, fig. 2), a well-defined
crust 1.25 to 1.5/u thick Is seen. Below this crust, the
wall Is constructed of irregular grains which tend to be
elongated in a direction normal to the wall surface.
Grains in the crust are smaller than those In the main
portion of the wall.
Islandlella lomltensls (Galloway and Wlssler)
Islandlella lomltensls (Galloway and Wlssler) has
both an internal toothplate and a radial wall and Is here
referred to this genus. The specimens were collected at a
depth of 205 m off the coast of southern California.
Test walls are about 30 /U thick and In thin section
under cross-polarized light have a very well-defined radial
extinction (pi. 40, figs. 4-5). Walls are fibrous in ap
pearance with a faint laminar structure. In tangential
section (pi. 40, fig. 6), the walls have a rather granular
appearance with a very diffuse pseudo-uniaxial cross figure.
PLATE 41
1-2 Islandlella callfornlca (Cushman and
Hughes)
1, surface of a late chamber. 2, cross
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure; C, secondary cal-
clte crust. X 20,000.
133
134
135
Pores on these specimens have an unusual appearance.
They are a series of parallel silts, each about 0.1 px wide
by 0.8/U long, and spaced 2 to 3/U apart (pi. 42, fig. 1).
Beside some of the pores are organic pore linings which
are only about 2 ja long, far too short to completely pene
trate the test wall. The Interpore regions are smooth,
marked by shallow grooves which delineate rhombic faces
0.05 to 0.1 /a across.
Cross-sections of the walls of this species have a
radiate fibrous structure (pi. 42, fig. 2). Grains are
elongated In a direction perpendicular to the wall sur
face. These grains are about 0.3M wide by up to 5/U
long. The outermost 2.5 /U of the wall consists of a layer
of shorter, less regular grains. This outermost layer Is
separated from the remainder of the wall by a thin Ir
regular band of organic matter.
Islandlella tortuosa (Cushman and Hughes)
Islandlella tortuosa (Cushman and Hughes) was re
ferred to this genus by N^rvang (1958)* However, no
internal toothplate could be observed by the present
author and no preferred extinctions were visible under
cross-polarized light. It Is possible that N/frvang's
observations were Incorrect and this species should remain
In the genus Cassldullna. The specimens were collected in
46 m depth of water off the coast of southern California.
PLATE 42
1-2 Islandlella lomltensls (Galloway and
Wlssler)
1, surface of a late chamber; P, pore;
PL, organic pore lining. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
136
137
138
Test walls are about 15 u thick and in both thin sections
and tangential sections under cross-polarized light, walls
are granular with no preferred extinctions (pi. 43, fig.
1) .
The surface of a late chamber Is shown In plate 44,
figure 1. Pores are thin slits 1.5 ja long by 0.1 yU wide
spaced 2 to 3 yU apart. These slits are not parallel, un
like the pores seen on I. lomltensls. Between pores, the
surface Is formed of flat calcite faces which vary In size
up to 0.5 /u.
In cross-section (pi. 44, fig. 2), the wall has a
curious herringbone structure. The majority of the grains
are plate-like, 1 to 2 yu long by 0.25 /u thick and oriented
at an angle to the test surface. Interspersed among these
flat grains are larger grains of irregular shape and up to
2 /u in diameter.
Florllus baslsplnatuB (Cushman and Moyer
Florllus baslBplnatus (Cushman and Moyer) is found
off the southern California coast at depths down to 300 m.
The walls of this species are about 6yU thick. Plate 43,
figure 2, shows a thin section under cross-polarized light
while plate 43, figure 3» is the same section rotated
through 45°. Although most of the wall is granular in
appearance, at some points a faint extinction can be seen
when the wall is positioned at 45° to the directions of
PLATE 43
All figures X 250; cross-polarized light
with polarizers oriented parallel to the
borders of the plate.
1 Islandlella tortuosa (Cushman and
Hughes)
Thin section.
2-4 Florllus baslsplnatus (Cushman and
Moyer)
2-3* thin section in two orientations
at 45° to show extinctions, indicated
by arrows, at 45° to the directions
of polarization. 4, tangential section.
5-6 Nonlonella mlocenlca Stella (Cushman
and Moyer)
1, thin section. 2, tangential
section.
139

PLATE 44
1-2 Islandlella tortuosa (Cushman and
Hughes)
1, surface of a late chamber; P, pore.
2, cross-section of the wall of a late
chamber; outer surface of the wall is
toward the right of the figure.
X 20,000.
141
142
143
polarization. Some of these points are Indicated on the
figures. In tangential section (pi. 43, fig. 4), no pre
ferred crystallographlc orientation Is detectable. Crystals
In the wall appear to be up to 2 u across.
Surface and cross-section views of late chambers of
specimens from a depth of 18 m are shown In plate 45.
Pores are about 1 yu in diameter, usually open, and with
organic pore linings beside them (pi. 45, fig. 1). They
are spaced 2.5 to 3*5 /u apart. The interpore regions are
formed of calclte faces measuring 0.1 /i In diameter or
smaller. Within restricted areas, these rhombic faces
have a constant orientation. These areas of constant
orientation are from 2 to 4 ja. across and have no well de
fined boundaries between them, but orientations in adjacent
areas may be quite different.
The test wall in cross-section (pi. 45, fig. 2) is
constructed of a series of laminae 1.5 to 3«0/u thick.
Each lamina is formed of a single layer of blocky grains
which extend across the thickness of the lamina and are
from 1 to 2 yu wide. On most of these grains, there are
rhombic faces which indicate that while crystallographlc
orientation is constant within grains, orientations of ad
jacent grains may be quite different.
The pores which are indicated on plate 45, figure 2,
do not penetrate the test wall completely. One pore passes
through the first two laminae and terminates at the
PLATE 45
1-2 Florllus baslsplnatus (Cushman and
Moyer)
1, surface of a late chamber of a speci
men from 18 m depth. 2, cross-section
of the wall of a late chamber of a sped
men from 18 m depth; outer surface of
the wall Is toward the right of the
figure; P, pores. X 20,000.
144

146
boundary of the third. Other poree begin and end within
the wall and appear to have no openings to either surface.
At a depth of 51 m, surfaces of late chambers have
a somewhat greater relief (pi. 46. fig. 1). Rhombic faces
are larger and less distinct. Some of the pores are
blocked or partially blocked by a secondary calclte crust.
The surface of a late chamber of a specimen from a
depth of 165 m is shown in plate 46, figure 2. Relief is
higher than on specimens from 51 m depth and most pores
are still open.
Plate 47 shows surface and cross-section views of
late chambers of specimens from a depth of 274 m, near the
lower end of the depth range for this species. Relief of
surfaces has decreased considerably (pi. 47, fig. 1) and
many pores are blocked by a secondary calclte crust. In
the interpore regions, very small calclte faces maintain a
constant orientation within restricted areas. These areas
are bounded by shallow sinuous grooves.
In cross-section (pi. 47, fig. 2), the laminae are
not as sharply defined as they were in specimens from
shallower water. Grains are larger and, like the
boundaries of the laminae, grain boundaries are less well
defined. A thin external layer, about 0.5/u thick, is the
secondary calclte crust.
As is the case with Bollvlna splssa. the crust on
F. baslsplnatus does not become well developed until near
PLATE 46
1-2 Florllus baslsplnatus (Cushman and
Moyer)
1, surface of a late chamber of a speci
men from 51 m depth. 2, surface of a
late chamber of a specimen from 165 m
depth. X 20,000.
147

PLATE 47
1-2 PlorlluB baslsplnatus (Cushman and
Moyer)
1, surface of a late chamber of a speci
men from 274 m depth; P, pore. 2,
cross-section of the wall of a late
chamber of a specimen from 274 m depth;
outer surface of the wall is toward the
right of the figure. X 20,000.
149

151
the lower depth limit for the specie8 although signs of
the crust are seen on specimens from shallower water. A
well developed crust on P. haslsplnatus. however, takes on
the pattern of sinuous grooves characteristic of a well
developed crust on Bollvlna argentea, rather than the zig
zag groove pattern seen on well developed crusts on B.
splssa.
Nonlonella mlocenlca Stella Cushman and Moyer
Like Florllus haslsplnatus, Nonlonella mlocenlca
Stella Cushman and Moyer Is a shallow water species found
off the coast of southern California at depths of less than
400 m. In thin sections and tangential sections, specimens
of N. mlocenlca Stella are very similar In appearance to
specimens of P. haslsplnatus except that the 45° extinc
tion Is not so noticeable on the former species (pi. 43,
figs. 5-6).
Plate 48 shows surface and cross-section views of
late chambers of specimens from 18 m depth of water. Pores
on this species are slightly smaller than those on P.
haslsplnatus, about 0.75yu In diameter (pi. 48, fig. 1).
Although most pores are open, many are blocked by a thin
secondary calcite crust. The interpore regions of the two
species are very similar, being formed of Irregular areas
2 to 3 p. across within each of which crystal orientations
are constant. Rhombic faces on N. mlocenlca Stella are not
PLATE 48
1-2 Nonlonella mlocenlca Stella Cushman and
Moyer
1, surface of a late chamber of a speci
men from 18 m depth. 2, cross-section
of the wall of a late chamber of a
specimen from 18 m depth; outer surface
of the wall is toward the right of the
figure. X 20,000.
152
153
2
154
as distinct as those on F. baslsplnatus.
In cross-section, specimens of both species from
this depth show strong similarities. The walls of N.
mlocenlca Stella are laminated, with laminae 1.5 to 2.0 yu
thick (pi. 48, fig. 2). Grains are 1 to 2/u wide, extend
ing across the full thicknesses of the laminae and oc
casionally transgressing laminar boundaries. Pores do not
penetrate the full thickness of the test wall.
Surfaces of specimens from a sample depth of 165 m
are considerably smoother (pi. 49, fig. 1) and most pores
are blocked by a secondary calclte crust. Organic pore
linings, each about 1 /a long, are seen lying on the surface
beside the pores.
At a sample depth of 274 m, flat calclte rhombs up
to 0.25 u wide may be seen on chamber surfaces (pi. 49,
fig. 2). There are no visible boundaries between larger
crystal units. Although pores are still visible, most are
blocked by secondary calclte crust.
Near the lower end of the depth range for this
species, at a depth of 366 m, the surfaces of late chambers
appear similar to surfaces of specimens from much shallower
water (pi. 50, fig. 1). Most pores are open and the inter
pore regions consist of Irregular areas 2 to 4 /a across
separated by sinuous grooves. Orientation of rhombic faces
is constant within each of these areas.
In cross-section, however, the test wall at this
PLATE 49
1-2 Nonlonella mlocenlca Stella Cushman and
Moyer
1, surface of a late chamber of a speci
men from 165 m depth. 2, surface of a
late chamber of a specimen from 274 m
depth. X 20,000.
155

PLATE 50
1-2 Nonlonella mlocenlca Stella Cushman and
Moyer
1, surface of a late chamber of a speci
men from 366 m depth. 2, cross-section
of the wall of a late chamber of a sped
men from 366 m depth; 0, outer surface
of the wall. X 20,000.
157

159
depth appears quite different (pi. 50, fig. 2). Laminae
are much thinner, from 0.25 to 1.5/1 thick. Grains within
each lamina are up to 3 p. wide, giving a brick wall ap
pearance to the chamber wall. Many adjacent grains show
the same crystallographlc orientation.
The development of a secondary calclte crust on F.
baslsplnatus and N. mlocenlca Stella Is not so apparently
connected with depth as It was on the two species of
Bollvlna, B. argentea and B. splssa. The crust on N.
mlocenlca Stella appeared thinner on specimens from the
deepest sample than It did on specimens from Intermediate
depths. It was noted that on the species of Bollvlna.
similar Inconsistencies occurred within limited portions
of the depth ranges of these species. Because of the much
greater depth ranges for Bollvlna. however, It was possible
to distinguish a regular development with depth.
In order to determine what effect, If any, fossill-
zatlon had on ultrastructures, a number of specimens of N.
mlocenlca Stella were taken from the Lower Pleistocene
sediments of Timms Point, California. Only those specimens
which showed very good preservation were selected. Surfaces
of late chambers of two of these specimens are shown In
plate 51. On one specimen (pi. 51* fig* 1)» there does not
appear to be a great deal of alteration. Pores are visible,
as is the secondary calclte crust, and organic pore linings
may be seen beside many of the pores. On the other spec!-
PLATE 51
1-2 Nonlonella mlocenlca Stella Cushman and
Moyer
1-2, surfaces of late chambers of two
different specimens from the Pleisto
cene of Timms Point. X 20,000.
160
161
2
162
men (pi. 51* fig* 2), however, the surface consists of
large Irregular calclte crystals, most over 1 p. in
diameter, which give the surface a configuration completely
unlike that on any of the Holocene specimens. These two
figures represent the two extremes of alteration which were
observed. Most specimens showed sufficient alteration that
the ultrastructures of the surface could not be easily com
pared with those of surfaces of Holocene specimens.
Cross-sections of Pleistocene specimens are shown
In plate 52. In neither cross-section Is the laminar
structure visible and grains are considerably larger than
In Holocene specimens. There Is little apparent relation
ship between the appearances of fossil and Holocene speci
mens.
No difference could be observed between Holocene
and fossil specimens In thin sections.
Cassldullna aff. C. laevigata d'Orblgny
A species collected at a depth of 420 m off the
west coast of South America Is here considered to have
strong affinities to Cassldullna laevigata d'Orblgny. The
specimens bear a close resemblance to d'Orblgny's model of
this somewhat problematical species.
Test walls are about 15p thick and show no pre
ferred crystallographic orientations In thin section under
cross-polarized light (pi. 53, fig. 1). There Is a
PLATE 52
1-2 Nonlonella mlocenlca Stella Cushman and
Moyer
1-2, cross-sections of the walls of late
chambers of two different specimens from
the Pleistocene of Timms Point; outer
surfaces of the walls are to the right
of the figures. X 20,000.
163

PLATE 53
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-2 Cassldullna aff. C. laevigata
d'Orbigny
1, thin section. 2, tangential
section.
3 Cassldullna carlnata Silvestri
Thin section.
4-5 Cassldullna pulohella d'Orbigny
1, thin section. 2, tangential
section.
165
166
3
167
granular appearance to the walls, the grains appearing to
be 2 to In diameter. In tangential section (pi. 53*
fig. 2), the same granular appearance Is observed.
The surface of a late chamber Is shown In plate 54,
figure 1. Pores are slightly under 2 /u In diameter and
are separated by distances of 3 to 6 p . Many of the pores
are blocked or partially blocked by secondary calcite
crust and organic pore linings are visible lying on the
surface. Interpore regions are formed of flat euhedral
calcite faces which do not maintain a constant orientation
aero88 the chamber surface.
In cross-section (pi. 54, fig. 2), the wall is con
structed of irregular grains averaging 0.5 in diameter.
There is no regularity to the arrangement of the grains
and considerable organic matter occurs between them. A
vague laminar structure parallels the surface. It was not
possible to observe any larger units which might be the
larger grains which seemed to be present when the tests
were observed In cross-polarized light.
Oaasldullna oarlnata Silvestri
Specimens of Cassldullna carlnata Silvestri were
obtained from the Pay of Biscay at a depth of 830 m. Walls
are from 5 to 6yu in thickness and have no preferred
crystallographlc orientation in thin section under cross
polarized light (pi. 53* fig. 3). In tangential sections,
PLATE 54
1-2 Cassldullna aff. C. laevigata d'Orbigny
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
168

170
walls appear to be very finely granular.
The available specimens had been glued to a faunal
slide with gum tragacanth. No amount of washing was able
to completely remove the gum from the tests and hence
surface replicas are somewhat blurred. The gum appears as
thin cloudy masses (pi. 55* fig. 1). Pores are about 1.5
yu In diameter, often surrounded by larger pore funnels.
Distances between pores are from 6 to 10^u. Interpore
regions are formed of smooth Indistinct calcite faces
averaging 0.1 yu In diameter.
In cross-section (pi. 55, fig. 2), a well defined
calcite crust 0.5 Ai thick Is separated from the remainder
of the wall by a layer of organic matter. The crust can
be seen to descend Into and coat the portion of a pore
funnel which Is visible In this figure. Grains are
generally less than 0.5yU In diameter, of Irregular shape
and arrangement, and with grain boundaries poorly defined.
Considerable organic matter occurs between grains.
Cassldullna pulohella d'Orblgny
Specimens of Cassldullna pulchella d'Orblgny were
taken from a sample collected of the west coast of South
America at a depth of 420 m. In thin section under cross-
polarized light, no preferred crystallographlc orientation
Is seen (pi. 53, fig. 4). Walls appear very finely granular
In tangential section (pi. 53» fig. 5)* The walls of this
PLATE 55
1-2 Cassldullna carlnata Silvestri
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure; PF, portion of a
pore funnel; C, secondary calcite crust.
X 20,000.
171

173
species are from 10 to 15 /i thick.
Plate 56 shows surface and cross-section views of
late chambers. Pores are 1.5 to 2.0^ In diameter* spaced
3 to 4 fi apart (pi. 56, fig. 1). Secondary calcite crust
partially blocks many of the pores and organic pore lin
ings can be seen beside many of these blocked pores. The
Interpore regions sure smooth and consist of very In
distinct calcite faces, most of which are under 0.1^ In
diameter.
In cross-section (pi. 56, fig. 2), there Is an Ir
regular poorly defined laminar structure with laminae 1.5
to 2.0 fa thick. Grains are of Irregular shape and grain
sizes vary from 0.5 to 4.0 fa. Grain boundaries are not
clearly defined.
Oassldullna brazlllensls Cushman
Cassldullna brazlllensls Cushman Is one of the most
thickly walled of the benthlc hyaline species with walls
often exceeding 70fa In thickness. The specimens were col
lected In the Antarctic at a depth of 3800 m.
In thin section under cross-polarized light, the
walls have a slightly fibrous appearance. There Is an
extinction when walls are oriented at 45° to the directions
of polarization and this extinction Is clearly shown In
plate 57, figures 1-2. The extinction appears as dark
bands 5 to 10/u wide which extend through the thickness of
PLATE 56
1-2 Caseldullna pulchella d'Orbigny
1, surface of a late chamber. 2t cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
174

PLATE 57
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-3 OaaBldullna brazlllensls Cushman
1-2, thin section In two orientations
at 45° to show the extinction at 45°
to the directions of polarization,
Indicated by arrows. 3, tangential
section.
4 Pullenla bulloldes (d'Orblgny)
Thin section; note extinction at 45°
to the directions of polarization,
lndloated by arrow.
5-6 Pullenla Salisbury! R. E. and K. C.
Stewart
5, thin section. 6, tangential
section.
176
177
0 - 1 mm.
178
the teat wall. A similar 45° extinction is seen on many of
the granular walled species. However, many of these species
are thin walled and this extinction is not usually as clear
ly visible as it is in C. brazlllensls. In tangential
section (pi. 57, fig. 3), no preferred extinctions can be
seen.
There is no sign of a secondary calclte crust on
either surface or cross-section views of late chambers
even though this species came from a very great depth.
Pores are open, often with organic pore linings beside
them (pi. 58, fig. 1), and distances between pores are in
excess of 10 u. Interpore regions are composed of smooth
flat calcite faces 0.5 to 2.0/1 across.
The wall in cross-section (pi. 58, fig. 2) closely
resembles the walls of C. carlnata with the exception that
no crust was seen on C. brazlllensls.
Pullenla bulloldes (d'Orbigny)
Specimens of Pullenla bulloldes (d'Orbigny) were
collected off the west coast of South America at a depth
of 2700 m. Walls of this species are about 25/i thick and,
in thin section under cross-polarized light, have a faint
extinction when walls are oriented at 45° to the directions
of polarization (pi. 57, fig* 4). There is a slightly
fibrous appearance but the wall appears generally granular,
particularly in the region of the proloculus. In tangential
PLATE 58
1-2 CasBldullna brazlllensls Cushman
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
179

181
section, there Is no preferred crystallographlc orientation
and grains appear to be 3 to 5 in diameter.
The pores on a late chamber (pi. 59, fig. 1) are
exceptionally small, averaging less than 0.25^ In diameter,
and are spaced 3 to 4^i apart. Interpore regions are
relatively smooth, although faint crystal faces may be
seen. A pattern of grooves outlines Irregular areas 2 to
7 /u across. These grooves have a rectangular configuration
which Is quite distinct from the sinuous and zigzag patterns
seen on other species.
An outer layer about 3 /U thick can be seen In cross-
section (pi. 59, fig. 2). This layer Is composed of very
large, flat grains up to 7/u long and of the same thickness
as the outer layer Itself. These grains correspond in size
to the areas on the surface delineated by the groove pat
tern.
Below the outer layer, the test is composed of ir
regular grains 0.5 to 2.0/u In diameter. Rhombic faces on
these grains indicate that adjacent grains do not have the
same crystallographlc orientation. Grains appear to be
bounded by layers of organic matter. There was no sign of
the fibrous structure which was seen in thin sections.
Pullenla sallsburyl R. E. and K. C. Stewart
A much smaller species of Pullenla. P. sallsburyl
R. E. and K. C. Stewart, was collected off the coast of
PLATE 59
1-2 Pullenla bulloldes (d'Orbigny)
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
182

184
southern California at a depth of 82 m.
Walls of this species are about 7 px thick and appear
granular in thin section under cross-polarized light (pi.
57» fig. 5). In tangential section (pi. 57, fig. 6), walls
appear to be formed of distinct grains up to 4 p. in
diameter.
Pores on this species are about the same size as
those on P. bulloldes but are much more closely spaced,
from 0.75 to 1.5/U apart (pi. 60, fig. 1). Interpore
regions are smooth, with a pattern of shallow grooves which
suggests the rectangular configuration seen on surfaces of
P. bulloldes.
In cross-section (pi. 60, fig. 2), the test is con
structed of irregular grains up to 1.5 /U in diameter.
There was a great deal of organic matter in the test,
particularly in the vicinity of pores. Numerous pores can
be seen to begin and end within the test wall and the
diameters of these pores are about twice the diameters of
pores as seen on the surface.
Pullenla malklnae Coryell and Mossman
Specimens of Pullenla malklnae were taken from a
sample collected at a depth of 420 m off the west coast of
South America.
Walls are about 25 /U thick and appear granular in
both thin sections and tangential sections under cross-
PLATE 60
1-2 Pullenla sallsburyl R. E. and K. C.
Stewart
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
top of the figure; P, pores. X 20,000.
185

187
polarized light (pi. 61, figs. 1-2). Grains appear to be
anywhere up to 5/i in diameter.
Surface and cross-section views of late chambers
are shown in plate 62. Pores average about 0.25^ in
diameter and the surface has the same rectangular groove
pattern as was seen on the other two species of Pullenla
(pi. 62, fig. 1). The areas enclosed by the grooves are 1
to 2^u across.
In cross-section (pi. 62, fig. 2), the walls are
constructed of irregular grains 0.5 to 1.0/a in diameter.
There is no regular arrangement to these grains and con
siderable organic matter occurs between them.
Chllostomella ovoldea Reuss
Chllostomella ovoldea Reuse may be considered as
typical of the granular walled group of hyaline fora-
mlnlfera, and was described and illustrated as such by both
Wood (19^9) and Towe and Cifelli (1967). The specimens
were collected in 632 m of water off the coast of southern
California.
Walls of this species are about 7 /u in thickness.
In thin section under cross-polarized light, they have a
characteristic granular appearance (pi. 61, fig. 3) with an
extinction at 45° to the directions of polarization. In
tangential section (pi. 61, fig. 4), the wall appears to be
made of large irregular grains up to 10/u long. These
PLATE 61
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-2 Pullenla malklnae Coryell and Mossman
1, thin section. 2, tangential
section.
5-4 Chllostomella ovoldea Reuss
1, thin section; arrows indicate ex
tinction at 45° to the directions of
polarization. 2, tangential
section.
188
189
2
i 0 1 m m »
4
PLATE 62
1-2 Pullenla malklnae Coryell and Moseman
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
190
191
192
grains are separated by very sharply defined, sinuous
boundaries. However, when the section Is rotated, Indivi
dual grains do not become completely extinct In any one
position. Rather, a portion of each grain may become ex
tinct while the remainder of the grain remains light.
Surfaces of two different specimens are shown In
plate 63, figures 1-2. Pores are not visible on either
figure though rarely a slit up to 3 yu long by O.lyu wide
may be seen. These slits may be pores reduced to slit-llke
openings by a secondary calclte crust. The surface shown
In plate 63, figure 1, Is smooth and Is divided into ir
regular areas by a pattern of sinuous grooves. These areas
are 1 to 6 yu in diameter and are probably the "grains'*
which are seen In tangential section under cross-polarized
light. The surface in plate 63, figure 2, Is composed of
angular grains averaging about 0.4/U across and separated
by a closer groove pattern.
In cross-section (pi. 63, fig. 3), there is a well
defined external layer 1.5 /u thick which Is composed of
Irregular grains averaging about 0.4/u In diameter. It is
the boundaries of these grains which may be seen on the
surface in plate 63, figure 2. Below this outer layer,
the test Is constructed of Irregular grains arranged in no
apparent pattern and varying In size up to 2 In diameter.
PLATE 63
1-3 Chllostomella ovoldea Reuss
1-2, surfaces of late chamber of two
different specimens. 3» cross-section
of the wall of a late chamber; outer
surface of the wall Is toward the right
of the figure, inner surface to the
left. X 20,000.
193

Gyroldlna neosoldanll Brotzen
195
The type species of the genus Gyroldlna. Gyroldlna
orbicularis d'Orbigny, was described by Wood (1949) as hav
ing a granular wall, and Loebllch and Tappan (1964) report
ed the genus as being granular walled. Specimens of
Gyroldlna neosoldanll Brotzen were collected In the Ant
arctic at a depth of 3800 m.
In thin section under cross-polarized light, walls
of this species have a distinct radial extinction (pi. 64,
figs. 1-2). Although this extinction Is not as complete as
Is the extinction on many other radial walled species, It
is nevertheless clearly visible, particularly on septa.
However, on tangential sections, or on fragments of crushed
tests (pi. 64, fig. 3), no preferred extinction is seen and
the test appears to be granular. The wall In thin section
has a fibrous, laminated structure.
On the surface of a late chamber (pi. 65, fig.l) are
pores about 0.5 u in diameter which are separated by dis
tances of 1.5 to 5*0 u. Most of the pores appear to be
open, filled with organic matter. Interpore regions are
smooth with faint calclte faces 0.25 to 0.5^i in diameter.
The cross section view (pi. 65* fig. 2) shows that
the wall Is constructed of irregular grains 0.25 to 1.0/u
in diameter, with grains generally becoming smaller toward
the outer surface. There is no regularity to the arrange
ment of grains and considerable organic matter occurs
PLATE 64
All figures X 250; cross-polarized light with
polarizers oriented parallel to the borders
of the plate.
1-3 Gyroldlna neosoldanll Brotzen
1-2, thin section in two orientations
to show the radial extinction. 3»
fragment of a test.
4-6 Hoeglundlna elegans (d'Orbigny)
4-5, thin section of two orientations
to show the radial extinction. 6,
tangential section showing a portion
of the pseudo-uniaxial cross figure
on one of the chambers.
196
197
3
i 0 - 1 mm. i
6
PLATE 65
1-2 Gyroldlna neosoldanli Brotzen
1, surface of a late chamber. 2, cross-
section of the wall of a late chamber;
outer surface of the wall Is toward the
right of the figure. X 20,000.
198
2
200
between them.
Hoeglundlna elegants (d'Orbigny)
Hoeglundlna elegane (d'Orbigny) Is the commonest
and most widely distributed of that small group of hyaline
foraminifera which secretes aragonltlc rather than calcltlc
tests. A recent Investigation by Reiss and Schneldermann
(1969) examined both surfaces and cross-sections of
Miocene and Recent specimens under the electron micro
scope. These authors reported that the wall was con
structed of hexagonal prisms of aragonite oriented with
the long axes of the prisms (and hence the crystallographlc
axes also) perpendicular to the wall surface. The wall
was laminated, with larger lamellae, formed by the suc
cessive secretion of chambers, divided Into smaller
"primary lamellae." Both types of lamellae were separated
from each other by layers of organic matter, while Indivi
dual prisms of aragonite were each sheathed In an organic
coating. The organic matter thus forms a series of layers
connected to each other by the grain sheaths to produce a
three dimensional reticulum. Secretion of mineral took
place within all of the "primary lamellae" of each of the
larger lamellae at one time.
The present author would urge caution In Inter
preting dark boundaries between grains as organic matter.
Grains, both on surfaces and in cross-sections, are usually
201
bounded by grooves, and replicas of such grooves may fold
over to give the appearance of lines of organic matter.
Some of the features Interpreted by Reiss and Schneldermann
as organic matter could well be such folds In the replica
films.
From a comparison of these authors' plates with
those taken during the present Investigation, It would ap
pear that a certain amount of alteration has occurred on
the Miocene specimens, though perhaps not as much as would
occur In smaller, calcltlc tests within the same time
Interval.
The specimens used In the present Investigation are
recent material collected off the west coast of South
America at sample depths of from 420 m to 2980 m.
In thin section under cross-polarized light, walls
have a strong radial extinction and a pronounced laminar
structure (pi. 64, figs. 4-5). In tangential section,
broad pseudo-unlaxlal cross figures may be seen on chambers
when the section Is rotated (pi. 64, fig. 6).
Pores occur on some areas of chambers and not on
others. It was not possible to distinguish any pattern to
the distribution of porous areas under the electron micro
scope because of the relatively large sizes of chambers,
nor, because of the very small size of the pores, could
such patterns be found by using light microscopy. The pos
sibility of an ordered arrangement to porous areas never-
202
theless still exists.
In plate 66, porous and pore-free regions on the
surface of a late chamber of a specimen from 420 m water
depth are shown. Pores are about 0.15/u In diameter and
are spaced at an average distance of 0.6/1 (pi. 66, fig.
1). Shapes of pores are somewhat angular In section.
Interpore regions are formed of rectangular grains averag
ing 0.1 fa wide by 0.4yu long. In the pore-free region,
the surface Is fluted, with flutlngs 0.25/U wide by 2.0/U
long (pi. 66, fig. 2). Sinuous grooves bound areas 4 to
6 /u across. Within each of these areas, fluting has a
constant direction. Flutlngs in adjacent areas may be In
either the same or In different directions.
At a sample depth of 735 m, the surface flutlngs are
considerably smaller— about 0.05/u by l.O/u (pi. 67# fig.
1). There are two major directions of fluting, oriented
at an angle of slightly under 65° to each other. Since the
angle between contact twins of aragonite is 63° 48', the
directions of fluting would represent orientations of such
contact twins. The flutlngs themselves are probably
lamellar polysynthetic twins. Angles between composition
planes of the contact twins and of the lamellar twins are
consistent with this Interpretation.
The contact twins produce a hexagonal pattern on
the surface, analogous to the pseudohexagonal prisms
produced by contact twins In megascopic crystals of
PLATE 66
1-2 Hoeglundlna elegans (d'Orbigny)
1, surface of a porous region on a late
chamber of a specimen from 420 m depth.
2, surface of a pore-free region on a
late chamber of a specimen from 420 m
depth. X 20,000.
203

PLATE 67
1-2 Hoeglundlna elegans (d'Orbigny)
1, surface of a late chamber of a
specimen from 735 m depth. 2, sur
face of a late chamber of a specimen
from 1130 m depth. X 20,000.
205

207
aragonite. Occasional grooves between these hexagonal
areas separate Individual grains. Note, however, that not
all grains are hexagonal in outline. These grains vary In
size from 0.5 to 2.0/1 across.
On the same figure, a distinct layer of greater
relief may be seen to overlie much of the surface. On
porous regions of the same specimen, pores occur both
within grains and along grain boundaries.
At a sample depth of 1130 m, flutings are larger,
0.1 /i wide by up to 2.0/i long, and relief is correspond
ingly greater (pi. 67, fig. 2). However, at a greater
depth yet, 2710 m, the flutings are smaller and much less
distinct (pi. 68, fig. 1). Groove patterns divide the
surface into areas averaging 2 /i across. Contact twins
occur In some, but not in all, grains.
Plate 68, figure 2, is a porous area on the surface
of a specimen from 2980 m depth. The flutings are even
less distinct at this depth, as are the grooves.
Cross-sections of H. elegans are shown in plate 69.
On a specimen from 420 m depth (pi. 69, fig. 1), two outer
laminae, each about 1 /i thick, are visible. In the outer
most lamina, grains are of irregular shape and arrangement,
averaging about 0.5/1 in diameter. The second lamina is
constructed of large flat grains 2 or more /a. in length
and extending across the thickness of the lamina. Below
these outer layers, the test is constructed of irregular
PLATE 68
1-2 Hoeglundlna elegans (d'Orbigny)
1, surface of a late chamber of a
specimen from 2710 m depth. 2, sur
face of a late chamber of a specimen
from 2980 m depth. X 20,000.
208

PLATE 69
1-2 Hoeglundlna elegans (d'Orbigny)
1, cross-section of the wall of a late
chamber of a specimen from 420 m depth;
outer surface of the wall is toward the
right of the figure. 2, cross-section
of the wall of a late chamber of a
specimen from 2980 m depth; outer sur
face of the wall is toward the right
of the figure. X 20,000.
210
2
212
grains, most under 1yU In diameter. Organic matter occurs
between grains and between laminae. In many places, the
organic matter produces a radial structure In the wall.
Plate 69» figure 2, Is a cross-section of the wall
of a late chamber of a specimen from a depth of 2980 m.
Indlctlnct laminae, 1.5 to 3.0yu thick, are constructed of
equant, blocky grains from 1 to 4 yu In diameter. Smaller
grains occur Interspersed among the larger ones. The
major change with depth Is an Increase In grain size and
an Increase in thickness of laminae.
Because of the laminated character of the test wall,
it was not possible to determine whether this species
develops a secondary crust or not. Although surface con
figurations change with Increasing depths, there are
similarities in all of these configurations and at no
depth do pores appear to be blocked. The layer overlying
the surface on plate 67» figure 1, does, however, suggest
secondary calcification.
Stomatorblna concentrlca (Parker and Jones)
A few specimens of another aragonltlc species,
Stomatorblna concentrlca (Parker and Jones), were obtained
from a sample collected In 100 m of water off Juan Fernan
dez Island. There were insufficient specimens for either
thin sections or cross-sections. However, a few surface
replicas were made and one of these is shown in plate 70.
Stomatorblna
Surface of a
PLATE 70
concentrlca (Parker and Jones)
late chamber. X 20,000.
213

Nowhere on these specimens was it possible to see
pores. Although Loeblich and Tappan (1964) reported the
genus as being aragonltlc, based on X-ray diffraction
studies, the surface Itself suggests calclte as the mineral
rather than aragonite, consisting as it does of a multitude
of rhombic faces. If sufficient material were available,
an investigation of the mineralogy of this species might
prove of interest since both minerals may be present.
216
ULTRASTRUCTURE OP BENTHIC HYALINE TESTS
Comparisons Between Light
and Electron Microscopy
The usual method of preparing thin sections of tests
Is by grinding. Such sections may range In thickness from
35 yU, the standard thickness for petrographlc sections,
down to as little as 5 yu, which Is about as thin as a
section can be ground without the use of special and
elaborate techniques. Grain sizes within any one test
seldom average greater than 2 yu in diameter. Hence, with
even the thinnest ground sections, observations are made
through a layer of superimposed grains. This has resulted
In some authors, e.g., Krashenlnnlkov (1956), Interpreting
crystallographlcally homogeneous regions In the test as
being single crystals. Towe and Clfelll (1967) discuss
this type of error In more detail and point to the diffi
culties of studying such fine structures by means of thin
sections.
One type of structure which is very frequently
observed under light microscopy Is a fibrous appearance of
the walls of some species. In many cases, no corresponding
structure can be observed under the electron microscope.
For example, the walls of Lentlcullna cultrata and
Gyroldlna neosoldanll are very fibrous in appearance In
217
thin section under either unpolarized or cross-polarized
light. Yet under the electron microscope, there Is no sign
of this fibrous structure. It Is possible that this
structure is produced by some three dimensional organiza
tion which, while visible in a section, is not apparent in
the planar replicas used in electron microscopy. Towe and
Cifelli (1967) attributed the fibrous appearance to the
superlmposltlon of pores in the section. However, this
seems unlikely since in many cases the structure is not
seen in the outer few microns of the test where pores are
still present, and it can be seen in very thin sections of
tests which do not have very porous walls.
Quite often, thin sections show a narrow band, up
to 10yu wide, on the outer surface of the wall (and usually
on the inner surface also). These bands lack the fibrous
structure seen on the remainder of the wall although they
have the same crystallographic orientation. In two species,
Marglnullna obesa and Lentloullna cultrata. this external
nonfibrous layer was clearly visible in thin sections. Yet
the structure of the outer parts of the wall, as seen under
the electron microscope, was not significantly different
from the remainder of the wall. If the phenomenon which
produces the fibrous structure were absent or were re
stricted in the outer few microns of the wall, this would
account for the presence of nonfibrous outer layers.
Grains and CrystalB
218
A b Towe and Cifelli (1967) pointed out, there is a
certain amount of ambiguity in the usages of the terms
"grain'* and "crystal." In this present paper, the term
"grain" has been restricted to discrete physical units
with more or less well defined boundaries, while the term
"crystal" is used for units which are sufficiently con
tinuous at the molecular level to produce a continuity of
crystal and cleavage faces. In tests of many of the
species examined, crystal or cleavage faces could be seen
on grains. Since these faces always have the same
orientations on the same grains, it is a reasonable as
sumption that these grains are monocrystalline.
Tests of four species, Bollvlna argentea. Bollvlna
splssa, Gancrls maoricus, and Hoeglundlna elegans. were
disaggregated in an ultrasonic generator and the resulting
particles were dispersed on a carbon film for examination
in the electron microscope. These particles were of ap
proximately the same sizes as the grains observed in
replicas of the test walls of the different species. Under
selected area diffraction, the particles had single
crystal patterns. It seems likely, then, the grains of
which tests are composed are in general monocrystalline.
It was not possible to determine tne exact crystal-
lographic orientation of grains in any of the calcltlc
species. Individual grains are too small to be observed
219
under the polarizing microscope, though preferred orienta
tions can be seen in many species. Information from
replicas was too scanty to permit the determination of
crystal orientation under the electron microscope, although
orientations of adjacent grains could sometimes be com
pared.
In the case of the aragonitic species, Hoeglundlna
elegans. twinning patterns on the surfaces of tests allowed
crystal orientation to be determined. In this species, the
c-axes of crystals are oriented normal to the wall surface
while the other crystal axes have no preferred orientations.
Since the optic axes and the c-axis for aragonite are very
close together, the walls have a radial extinction.
Organic Matter
Organic matter occurs within the tests in patches
disseminated among grains, in discontinuous layers between
laminae, and, in some species, as linings to pores. As was
pointed out above, under the description of Hoeglundlna
elegans. care must be taken in interpreting dark regions on
replicas as organic matter. Seep grooves, folds in the
film, or even plastic from a primary replica can be con
fused with such material. That considerable organic matter
does occur in tests is readily apparent from observations
when tests are dissolved in dilute acids. This organic
matter forms the matrix, discussed by Towe and Cifelli
220
(1967)* within which calcification takes place. It occurs
in most species as a three dimensional retioulum with most
of the strands perpendicular to the wall surface. In those
species in which the structure of the wall has a more de
finite pattern, the matrix occurs as a series of discon
tinuous layers or two dimensional nets, parallel to the
wall surface and connected by a more random pattern of
strands.
It is suggested that these organic reticula produce
the fibrous appearance seen in thin sections of some tests.
Restriction of the reticulum to the central regions of the
wall would produce the nonfibrous outer layers. On
replicas, such a reticulum would appear as Isolated
patches of organic matter rather than as a series of
strands.
Pores
On many species, a secondary calclte crust blocks
pores completely or else reduces them to thin slit-like
openings. In other species, pores apparently began and
ended within the walls. Pores which are blocked, or which
do not penetrate the wall completely, could not provide any
sort of communication between external and Internal portions
of the cell, and lead to the question of what functions the
pores serve. Because of the organic pore linings and the
organic sieve plates which are sometimes observed within
221
pores, It Is not likely that they are non-functlonal.
Since calcification Is most probably extracellular, It
would be necessary for the cell to be withdrawn from the
matrix during the calcification process. It is suggested
that pores enclose projections of the cell wall which may
werve as anchors to the matrix while at the same time
holding the cell wall away from the sites of calcification.
Laminae
The term “laminae" was used In the present paper In
reference to the thin layers, usually less than a micron
thick, of which many tests are constructed. These layers
must not be confused with the much thicker "lamellae"
which are observed In thin sections and which appear to be
connected with the secretion of successive chambers.
Lamellae are difficult to observe and trace under the
electron microscope because of their relatively large
sizes, from 5 to lOyu In thickness. The laminae are ap
parently the same structures as the "primary lamellae" dis
cussed by Reiss and Schneldermann (1969)*
Laminar structure In test walls seems to be deter
mined by the form of the organic matrix since organic
matter tends to be concentrated along the laminar
boundaries.
The Secondary Calcite Crust
222
A certain amount of secondary calcite crust was ob-
\
served on almost all of the specimens examined in this
investigation. The rajor exception was on those specimens
of Bollvlna argentea which were collected in a low oxygen
environment. On most of these specimens, the entire sur
face of the test was extremely smooth with no signs of
crustal development.
Development of the crust proceeds in a series of
stages:
1. At its inception, the crust appears as a thin
layer of very small calcite crystals, averaging less than
0.1^ in diameter, and coating the entire chamber surface,
including the surfaces of pore funnels. On radial walled
species, these crystals may vary in orientation across the
chamber surface, with no sharp boundaries between regions
of different orientation. On granular walled species,
orientation is consistent within restricted areas which are
separated from each other by more or less well defined
grooves.
2. The second stage is the development of a zigzag
groove pattern which delineates angular areas averaging
about 1 pi across. The surfaces of these angular areas are
smooth and their shapes suggest a mass of euhedral calcite
crystals. This stage is common on radial walled species,
but is seldom well developed on those species in which the
223
walls are granular.
3. The third etage Is the development of a
relatively high relief In which any pattern of the con
figuration of the test surface Is difficult to observe.
4. The final stage Is the development of a pattern
of sinuous grooves which delineates smooth areas, Irregu
lar In outline and several microns across. In many cases,
rhombic faces can be seen on these smooth areas within
each of which they maintain a constant orientation. These
areas are developed by coalescence of the smaller angular
areas seen In the second stage and the sinuous grooves are
developments from a zigzag groove pattern.
Although, In general, the successive stages develop
within a species with Increasing depths, correlation be
tween crustal development and depth Is by no means exact.
Many specimens will show stages that one would expect to
find on specimens from different depths. It will be
remembered that lack of oxygen can Inhibit crustal develop
ment and It Is entirely possible that other environmental
factors may also exert some control.
The stage of crustal development reached at any one
depth and the extent to which the crust will develop are
characteristics peculiar to the Individual species.
Since the crust Is seldom more than 1 px thick, It Is
unlikely to be observed In thin section because of the
Interference at the Interface between the test wall and the
224
embedding medium. Efforts to observe the crust on speci
mens of Bollvlna argentea were unsuccessful even when
specimens were used in which the crust was well developed.
The crust cannot be observed on whole specimens under the
light microscope. Although a well developed crust may com
pletely block pore openings, the pores remain as structures
in the wall and are still visible. The only case in which
the crust could be observed under the light microscope was
in tangential sections of Chllostomella ovoldea where the
' ‘grains” observed appeared to be the result of development
of a sinuous groove pattern on the surface.
The secondary calcite crust is not related to the
nonfibrous outer band which is visible in thin sections
and which was discussed above. This band does not differ
greatly in structure from the remainder of the test wall
as does the crust. It does not block pores and it does not
produce the characteristic surface configurations of the
crust. It may be up to 10/u thick, whereas the crust is
seldom more than 1.5/U thick. On many species in which a
well developed crust is present, no nonfibrous outer layer
is visible, and on other species in which the crust is only
slightly developed, there is a very clear outer layer.
The structure of the crust is similar in all species,
appearing as a single layer of large tabular grains, the
boundaries of which are generally not clearly defined.
Radial versus Granular Walls
225
In thin sections of many of the granular walled
species, an extinction could be observed when walls were
positioned at 45° to the directions of polarization. The
extinction occurs In only a small portion of the wall, un
like the extinction in radial walled species, and appears
to Involve clumps of grains of similar crystallographlc
orientation. This tends to confirm Towe and Cifelli's
(1967) statement that in granular walled tests, crystals
are oriented with the (1011) planes parallel to the wall
surface. Any such crystals in which the c-axes were in the
plane of the section would become extinct when the surface
was at an angle of 45° 22' to the directions of polariza
tion, this being the angle between the c-axls and the
(1011) planes in calcite.
Extinction in the radial walled species Is not con
fined to the portion of the wall which Is precisely parallel
to the directions of polarization. Rather, it may Involve
portions of the wall which are as much as 5° from parallel
ism. This Indicates that crystal orientation in these
species is not perfectly normal to the wall surface but may
vary through as much as 10°.
There is no apparent correlation between the optical
extinction of the wall and the structure of the wall. Cer
tain species, such as Lentlcullna cultrata. have a very
strong radial extinction, yet have walls constructed of
226
Irregular grains arranged In no apparent pattern. Many of
the granular walled species have a very regular arrangement
to the grains.
There also does not appear to be any correlation be
tween optical properties and other ultrastructural features
such as pore spaclngs, grain size and so forth.
One group of foraminifera, exemplified by Gyroldlna
neosoldanll, has walls Intermediate In optical properties
between radial and granular. These walls do not have a
preferred extinction In tangential sections nor on frag
ments of crushed tests. In thin sections, however, a
radial extinction is readily apparent. Towe and Cifelli
(1967) report a similar crystallographlc arrangement to
Clblcldes refulgens and described this type of wall as
Mindistinctly radial." Since a similar term had been used
by Krasheninnikov (1956) to describe a different type of
wall, the present author prefers to designate this group as
"intermediate.” Walls in this group are composed of grains
in which either the preference of orientation is slight,
or in which both orientations occur.
The Process of Calcification
A model for the calcification process in the fora
minifera was proposed by Towe and Cifelli (1967) who sum
marized and added to the available knowledge on the sub
ject. Calcification is most probably extracellular and
227
begins with the secretion of an organic matrix which con
sists of two components— a passive phase which gives shape
and form to the chamber to be secreted, and an active phase
which causes precipitation. A solution of the appropriate
ions is secreted into this matrix and these ions precipi
tate at the sites of the active phase.
The following additional comments are based on ob
servations made during the course of the present investiga
tion. Since the organic matrix is secreted extracellularly,
it must be formed within an invagination of the cell wall.
It is most likely in the form of a three dimensional
reticulum of passive phase, with lesser amounts of active
phase at the points where crystal growth is to be initiated.
Short extensions of the cell wall project into this
reticulum, possibly to anchor it to the cell wall. Once
calcification of a chamber is completed, these projections
may be withdrawn or become atrophied, leaving voids in the
test wall which are the pores. In some species, calcifica
tion takes place in a series of stages, since abandoned
pores may be seen within the wall with no connection to
either surface.
The fibrous structure visible in thin sections of
so many tests is most likely produced by the presence of
remnants of the passive phase. Once the chamber has taken
shape, calcification can proceed without the further neces
sity of the passive phase, using only the active phase and
228
the mineralizing solution. Thus, the passive phase, and
consequently the fibrous structure of the test wall, would
be absent from the outer few microns of the wall and pro
duce the nonfibrous outer layers.
Secretion of the mineralizing soluti n may continue
at all times, at rates which vary with the species in
volved, with environmental factors, and with the stage of
chamber development. Since the external portions of the
cell extend down from the aperture to cover early chambers,
this continuous secretion supplies the ions from which the
secondary calcite crust is formed. The pre-existing cal
cite of the early chambers can act as an active phase to
cause precipitation. The crust would be thickest on the
early chambers which are exposed to the mineralizing
solution for the greatest length of time and the con
tinuity of secretion would account for the relatively
homogeneous structure of the crust.
Although resorption appears to be an important
process in many calcifying organisms, no evidence of re
sorption was seen during this investigation.
229
ENVIRONMENTAL CONTROL OP ULTRASTRUCTURES
The most obvious changes In ultrastructure which are
produced by changes In environment are in the development
of the secondary calcite crust. This crust becomes thicker
and changes In Its appearances with Increases In depth. As
was noted under Bollvlna argentea. oxygen concentration
also affects the development of the crust. Other environ
mental factors such as pressure or pH may be involved as
well.
The degree to which the test wall is laminated is
depth controlled In many species. In both Bollvlna
argentea and Nonlonella mlocenlca Stella, laminae become
more sharply defined with increasing depths, and as a re
sult of this, grains become smaller and more regular in
shape and arrangement. Similar changes occur in Cancrls
maorlcus, though to a much lesser degree. In Florllus
baslsplnatus. however, lamination is less apparent at
greater depths and grain boundaries are less sharply
defined, while in Bollvlna splssa. little change could be
noted with changes in depth. Laminae in the aragonitic
species Hoeglundlna elegans become thicker with Increasing
depths and grains become larger and of more regular shape.
As is the case with the crust, other environmental factors
may also exert some control on the development of laminae.
In general, those species which live in deeper water
230
have more widely spaced pores than do congeneric species
from shallow water, even though pores may be otherwise
similar.
Ultrastructures are probably not of any great use
for paleoecological investigations. Considerable changes
can be produced by recrystallization within a relatively
short period of time, making it difficult to compare
fossil with Recent material. Furthermore, sensitivity of
ultrastructures to environmental changes is not very great
and the relationships between the two may be very complex.
It seems likely, then, that other methods of investigation
would be Just as accurate if not more so, and would pro
bably be more rapid.
231
ULTRASTRUCTURES AND GENETIC RELATIONSHIPS
As was shown in the study of Bollvlna argentea.
environmental factors can so radically alter the ultra-
structures of the test that comparisons of related species
may be difficult unless samples from a variety of environ
ments are used. Bollvlna argentea and Bollvlna splssa have
very similar depth and geographical distributions. Samples
of these two species came from the same geographical region
and from almost the same depth range. In many cases, speci
mens of the two species were taken from the same sample.
Yet it is difficult to see any strong resemblances between
the ultrastructures of the two species. The secondary
calcite crust becomes well developed in B. argentea but is
scarcely visible on B. splssa except on specimens from the
lower end of the depth range. The pores on B. splssa are
much larger and lack the conspicuous pore funnels observed
on pores of B. argentea. In cross-sections, the grains in
the walls of B. argentea are smaller and of less regular
siape than those of B. splssa. while the laminae which
develop in the walls of B. argentea are poorly developed in
&. splssa. The only structures in common between the two
species are the wall thicknesses and the spacings between
pores, which are comparable, and the radial extinctions,
which are similar.
Only one sample was used for each of the other five
232
species of Bollvlna which were examined. This sample would
represent only a limited portion of the environmental range
for that species. However, If genetic relationships had
produced ultrastructural similarities, each of these
species should have been comparable to at least one of the
samples of either B. argentea or B. splssa. Such was not
the case. Every one of the seven species appeared to have
Its own peculiar ultrastructure which bore little
resemblance to those of any of the others. The only
features In common were the radial extinctions and, In
most cases, the pore spacings.
Of the three species of Islandlella which were
examined, I. tortuosa had an ultrastructure quite dif
ferent from that of the other two species. As was noted,
this species should probably be retained in the genus
Oassldullna; it bore, however, little resemblance to any of
the species of Oassldullna either. I. callfornlca and I..
lomltensls appeared somewhat similar to each other in cross-
section, although coming from different depths and dif
ferent geographical regions.
There were remarkable similarities among the four
species of Oassldullna. Pores are comparable in size and
shape, and there are strong resemblances in the cross-
section views (cf. pis• 14, 15, 16, 18). The species
which came from the deepest water, 0. brazlllensls. dif
fered from the others in having greater distances between
233
pores.
The three species of Pullenla also resembled each
other closely. Pores were comparable in size and shape,
although pore spacing was by far greatest on Pullenia
bulloides. the species from the deepest water. All three
species had a characteristic groove pattern on chamber
surfaces in which angles of grooves are close to 90°,
producing a rectangular configuration to the pattern. In
cross-sections, walls of all three species are laminated,
and grains are large and blocky. Towe and Cifelli (1967)
illustrated surface and cross-section views of Pullenla
qulnqueloba (Reuss). The cross-section view resembles
those of the three species examined in the present investi
gation, although the surface lacks the rectangular groove
pattern and pores are much larger.
Hay, et al. (1963) illustrated a surface view of
Lentlcullna midwayensis (Plummer), and Towe and Cifelli
(1967) illustrated both surface and cross-section views of
Lentlcullna calcar (Linne). It is thus possible to make
comparisons between those species and Lentlcullna cultrata.
The surface views of L. calcar and L. cultrata are
similar, though pores on the latter species are about twice
the diameter as those on the former. Pores on L. midway-
ensiB are muoh larger yet, and of a much more regular
shape. The cross-sections of L. calcar and L. cultrata
show few similarities.
234
The two species which bore the greatest resemblances
to each other were Florilus basispinatus and Nonionella
miocenica stella. Both in turn showed strong similarities
to the species of Pullenia and to the figures of Nonlo-
nelllna labradorica (Dawson) given by Towe and Cifelli
(1967). Florilus, Nonionella, Pullenla and Nonlonelllna
are four closely related genera. In all four genera,
tests are laminated and constructed of large, blocky
grains, while pores tend to be small, averaging about
0.25 u in diameter.
In general, ultrastructure is consistent within
species, the only variations being those imposed by
environmental factors. The majority of specimens of a
single species taken from the same sample will show only
a slight amount of variation.
The optical properties of test walls are also con
sistent within species, and with the single exception of
Islandlella tortuosa, were found to be consistent within
genera. It is believed that a reinvestigation of the
genera Islandlella and Oassldullna might resolve this
particular anomaly.
Those genera, such as Clblcldes and Gyroldlna, in
which the wall is of an intermediate type between radial
and granular, also deserve further attention. The inter
mediate wall cannot be defined except by means of thin
sections, and it is possible that certain genera which
235
have been described as granular on the basis of an
examination of a fragment of a test, may actually have
Intermediate walls. The Intermediate wall type suggests
that a gradation may exist between the radial and granular
groups. If this Is so, the use of optical properties for
describing taxa would involve more complete descriptions
of the wall as seen in thin section under cross-polarized
light, rather than a simple division Into radial, Inter
mediate and granular. It is possible that subdivisions of
these groups, similar to those suggested by Krasheninnikov
(1956), may be feasible.
If due allowance is made for the influence of
environmental factors, ultrastructural characteristics can
be used in the definition of a particular species. This is
particularly well shown in the cases of Bollvlna argentea
and B. aenarlensls which are difficult to distinguish on
the basis of the gross morphologies of the tests alone, but
which had striking differences in ultrastructures. How
ever, the possibility exists of true sibling species in
which both gross morphologies and ultrastructures are the
same.
In the case of taxonomic units above the species
level, ultrastructure may indicate possible relationships.
However, the absence of similar ultrastructures does not
argue a lack of relationship, as witness the seven species
of Bollvlna, all of which had different ultrastructures.
236
The most useful distinguishing characteristics appear to be
pore size and shape, the structure of the wall as seen In
cross-section, and the optical properties of the wall.
Least useful characteristics are surface configurations and
distances between pores.
Ultrastructural studies can thus be used for
systematic purposes as an adjunct to the more classical
methods which utilize gross morphologies and lineages. The
use of ultrastructures would be generally limited to Recent
material because of the changes wrought by recrystallization
in fossil specimens.
237
SUMMARY AND CONCLUSIONS
Tests of benthic hyaline foraminifera are con
structed of monocrystalline grains which vary In size from
1/2 to 4/u In diameter. Observations tend to confirm Towe
and Clfelli's (1967) description of crystallographic
orientations. In radial walled tests, grains have a pre
ferred orientation In which the (0001) planes of crystals
are parallel to the wall surface; in granular walled tests,
the (1011) planes of crystals have a preferred orientation
parallel to the wall surface. There is a group of hyaline
foraminifera, which includes such genera as Clblcldes and
Gyroldlna. in which the wall is of an intermediate type
between radial and granular. In walls of this group,
either both types of orientation occur, or else the pre
ference of orientation is only slight. This type of wall
can be defined only by means of thin sections, and not by
means of tangential sections or by using fragments of
crushed tests. The type of wall is consistent within
species and probably within genera and higher taxa.
Anomalies appear to be due to faulty observations.
The postulated calcification process begins with the
extracellular secretion of an organic matrix. This matrix
occurs as a three dimensional reticulum which may be either
a series of irregular strands which are generally perpendi-
cular to the wall surface, or a series of layers, parallel
to the wall surface and connected to each other by Irregu
lar strands. The presence of the former type of matrix
produces the fibrous appearance seen In walls of some tests
In thin section; the latter type of matrix causes the
development of laminae within the wall. Secretion of the
matrix is followed by secretion of a mineralizing solution
from which calcium carbonate is precipitated into the
matrix. Precipitation may continue after the matrix it
self has become completely mineralized and thus produce
the nonfibrous outer layers which are seen in some tests
in thin sections. Projections of the cell wall extend into
the matrix, possibly to anchor the matrix to the cell wall.
These projections leave voids, which are the pores, in the
test wall after calcification has been completed. Secre
tion of each chamber wall proceeds in a series of stages in
some species. A continuous secretion of the mineralizing
solution between the cell wall and the test wall causes the
deposition of the secondary calclte crust.
Environmental factors control the extent to which
the secondary calclte crust develops in each species. In
most species, the degree to which the test wall is laminated
is also controlled by environment. The degree of lamina
tion in turn determines the size, shape and arrangement of
grains. The secondary calclte crust becomes thicker with
increasing depth and, as it increases in thickness, produces
239
a series of characteristic surface configurations. Lower
ing of oxygen concentrations inhibits the development of
the crust. Other environmental factors may also exert some
control. The degree of lamination may either increase or
decrease with increasing depths; in some species, there is
little change in the degree of lamination with changes in
depth. An Increase in the degree of lamination causes
grains to be smaller and more regular in shape and arrange
ment.
Among congeneric species, pores appear to be more
widely spaced on species which live at greater depths than
on species from shallower water.
Test ultrastructure is consistent within a species
except for changes which can be attributed to environmental
factors. There is little variation among specimens of the
same species taken from the same environment. Congeneric
species, even those in which gross morphologies are very
similar, may have very different ultrastructural features.
Absence of ultrastructural resemblances does not, there
fore, necessarily mean an absence of close genetic relation
ship. Within some genera, on the other hand, ultrastruc
tures of different species may be very similar. Re
semblances may also occur between ultrastructures of species
from different but related genera. Hence, similarities in
ultrastructures, in conjunction with other factors such as
similarities in gross morphologies, distributions, and
240
lineages, may be used to indicate relationships. The most
useful ultrastructural features for systematic purposes are
the size and shape of pores, and the appearance of walls in
cross-sections. The least useful ultrastructural feature
is the surface configuration, which is controlled mainly
by environmental factors.
Possilization may produce extreme and erratic
changes in ultrastructure because of recrystallization.
Extreme caution would be necessary in any attempts to use
ultrastructural features for paleoecological investiga
tions or for systematic studies among fossil species.
REFERENCES
241
FAUNAL REFERENCE LIST
Faunal index slides of the species which were
examined are on file in the Allan Hancock Foundation of the
University of Southern California. In the following list,
reference is made to publications in which these species
have been accurately illustrated.
Bollvlna argentea Cushman: Lutze (1964), text-
figure 9.
Bollvlna splssa Cushman: Lutze (1962), plate 24,
figures 14-18.
Bollvlna acuminata Natland: Bandy (1953), plate 24,
figure 6.
Bollvlna aenarlensls (Costa): Cushman (1937)»
plate 12, figure 26.
Bollvlna lnter.luncta Cushman: Cushman (1937),
plate 15, figures 1-3.
Bollvlna pseudobevrlchl Cushman: Cushman (1937),
plate 19, figure 5»
Bollvlna semlnuda Cushman: Cushman (1937), plate
18, figure 14.
Cancrls maorlcus Finlay: Finlay (1940), plate 64,
figures 102-104.
242
243
Lentlcullna cultrata (Montfort): Cushman and
McCulloch (1950), plate 37» figures 5-6.
Marglnullna obesa Cushman: Barker (I960), plate 65,
figures 5-6.
Islandlella callfornica (Cushman and Hughes):
Bandy (1953), plate 25, figure 1.
Islandlella lomltensls (Galloway and Wissler):
Bandy (1953), plate 25, figure 5«
Islandlella tortuosa (Cushman and Hughes): Bandy
(1953), plate 25, figure 3.
Plorllus baslsplnatus (Cushman and Moyer): Bandy
(1953), plate 21, figure 13.
Nonlonella mlocenlca stella Cushman and Moyer:
Bandy (1953), plate 22, figure 2.
Cassldullna aff. C. laevigata d'Orbigny: refer to
d'Orbigny's model; also Loeblich and Tappan (1964), text-
figure 604, figure 1.
Cassldullna carlnata Sllvestrl: Silvestri (1896),
plate 2, figure 10.
Cassldullna pulchella d'Orbigny: d'Orbigny (1839)
plate 8, figures 1-3.
Cassldullna brazlllensls Cushman: Cushman (1922),
plate 25, figures 4-5.
Pullenla bulloldes (d'Orbigny): Barker (I960),
plate 84, figures 12-13.
Pullenla sallsburyl R. E. and K. C. Stewart: Cush
man and Laiming (1931), plate 14, figure 2.
Pullenla malklnae Coryell and Mossman: Coryell and
Mossman (1942), plate 36, figures 3-4.
Chllostomella ovoldea Reuse: Cushman (1924), plate
1, figure 1.
Gyroldlna neosoldanll Brotzen: Barker (I960), plate
107, figures 6-7.
Hoeglundlna elegans (d'Orbigny): Bandy (1953)*
plate 23, figure 9.
Stomatorblna concentrlca (Parker and Jones):
Barker (i960), plate 105, figure 1.
0
REFERENCES
Bandy, 0. L., 1953. Ecology and paleoecology of some
California foraminifera. Part 1. The frequency dis
tribution of Recent foraminifera off California. Jour.
Paleo., vol. 27, no. 2, p. 161-182, pis. 21-25, text-
figs. 1-4.
Barker, R. W., I960. Taxonomic notes on the species figur
ed by H. B. Brady in his report on the foraminifera
dredged by H. M. S. Challenger during the years 1873-
1876 accompanied by a reproduction of Brady's plates.
Soc. Econ. Paleo. Miner., Spec. Pub. 9, p. i-xxlv,
1-238, pis. 1-115.
Bartlett, G. A., 1967. Scanning electron microscope:
potentials in the morphology of microorganisms.
Science, vol. 158, no. 3806, p. 1318-1319, text-figs.
1-2.
Be, A. W. H., 1965. The Influence of depth on shell
growth in Globlgerlnoldes saccullfer (Brady). Micro-
paleontology, vol. 11, no. 1, p. 81-97, pis. 1-2.
Be', A. W. H., McIntyre, A., and Breger, Bee L., 1966.
Shell microstructure of a planktonic foraminifera,
Globorotalla menardll (d'Orbigny). Eclogae Geol.
Helv., vol. 59, no. 2, p. 885-896, pis. 1-17, text-
figs. 1-2.
Coryell, H. N. and Mossman, R. W., 1942. Foraminifera from
the Charco Azul formation, Pliocene, of Panama. Jour.
Paleo., vol. lo, no. 2, p. 233-246, pi. 36.
Cushman, J. A., 1922. The foraminifera of the Atlantic
Ocean. U. S. Nat. Mus., Bull. 104, Pt. 3, Textu-
lariidae, p. i-viii, 1-149, pis. 1-26.
_______ , 1924. The foraminifera of the Atlantic Ocean.
U. S. Nat. Mus., Bull. 104, Pt. 5, Chllostomellldae
and Globlgerlnldae. p. i-v, 1-55, pis. 1-8.
245
246
Cushman, J. A., 1936. A monograph of the Bubfamlly
Vlrgullnlnae of the foraminiferal family Bullmlnldae.
Cushman Lab. Poram. Res., Spec. Pub. 9» p. i-xv, 1-228,
pis. 1-24.
Cushman, J. A. and Lalmlng, B., 1931. Miocene foraminifera
from Los Sauces Creek, Ventura County, California.
Jour. Paleo., vol. 5, no. 2, p. 79-120, pis. 9-14,
text-figs. 1-5.
Cushman, J. A. and McCulloch, Irene, 1950. Some Lagenldae
in the collections of the Allan Hancock Foundation.
Allan Hancock Pao. Exped., vol. 6, no. 6, p. 295-364,
pis. 37-48.
Finlay, H. J., 1940. New Zealand foraminifera; key
species In stratigraphy--no. 4. Roy. Soc. New Zealand,
Trans. Proc., vol. 69, pt. 4, p. 448-472, pis. 62-67.
Hansen, H. J., 1967. A technique for depiction of grind
sections of foraminifera by aid of compiled electron-
mlcrographs. Dansk Geol. Foren., Medd., vol. 17, no.
1, p. 128-130, pis. 1-2.
Hay, W. W. and Sandberg, P. A., 1967. The scanning
electron microscope, a major break-through for micro
paleontology. Micropaleontology, vol. 13» no. 4,
p. 407-418, pis. 1-2.
Hay, W. W., Towe, K. M., and Wright, R. C., 1963. Ultra
microstructure of some selected foraminiferal tests.
Micropaleontology, vol. 9» no. 2, p. 171-195, pis.
1-16.
Honjo, S. and Berggren, W. A., 1967. Scanning electron
microscope studies of planktonlc foraminifera. Micro-
paleontology, vol. 13, no. 4, p. 393-406, pis. 1-4,
text-figs. 1-2.
Hyde, P. and Krinsley, D., 1964. An improved technique
for electron microscopic examination of foraminifera.
Micropaleontology, vol. 10, no. 4, p. 491-493, text-
figs. 1-3.
Jahn, Brigitte, 1953* Elektronen mikroskoplsche Unter-
suchungen an Foramlniferenschalen. Zeltschr.
Wissensch. Mikroskopie, vol. 61, no. 5» p. 294-297,
text-figs. 1-9.
247
Krasheninnikov, V. A., 1956. Mikrostruktura stenkl
nekotorikh Kaenozoesklkh foramlnlfer 1 metodika ee
lzuchenlya v polarizovannom Bvete. Voprosy Mikro-
paleontologii, no. 1, Akad. Nuak SSSR, Otdel Geol.-
Geog. Inst., p. 37-48, pis. 1-2, text-flg. 1.
Krinsley, D. and Be*, A. W. H., 1965. Electron microscopy
of Internal structures of foraminifera. In: Kummel,
B. and Raup, D., eds., Handbook of paleontological
techniques. San Francisco: W. H. Freeman and Company,
p. 335-343, text-figs. 1-3.
Loeblich, A. R., Jr., Tappan, Helen, et al., 1964.
Treatise on invertebrate paleontology (Ed., R. C.
MOore), Part C: Sarcodlna chiefly "Thecamoebians"
and ForamlnlferIda. New York: Geol. Soc. America,
vols. 1-2, p. i-xxxl, 1-900, text-figs. 1-653.
Lutze, G. F., 1962. Variationsstatistik und Okologie bei
rezenten Foraminiferen. Pal. Zeitschr., vol. 36,
nos. 3-4, p. 252-264, pi. 24, text-figs. 1-3.
_______ , 1964. Statistical investigations on the
variability of Bollvlna argentea Cushman. Cushman
Found. Foram. Res. Contr., vol. 15, pt. 3, p. 105-116,
text-figs. 1-9.
Lynts, G. W. and Pfister, R. M., 1967. Surface ultra
structure of some tests of Recent Foramlnlferlda from
the Dry Tortugas, Florida. Jour. Protozool., vol. 14,
no. 3, p. 387-399, text-figs. 1-16.
N^rvang, A., 1958. Islandlella n. £. and Cassldullna
d'Orbigny. Vldensk. Medd. Dansk Naturhlst. Foren.,
vol. 120, p. 25-41, pis. 6-9.
/
d'Orbigny, A. D., 1839. Voyage dans l'Amerique Meridionale;
Foraminlferes. P. Bertrand, Strasbourg, vol. 5, P.
1-86, pis. 1-9.
Orr, W. N., 1967. Secondary calcification in the fora
miniferal genus Globorotalla, Science, vol. 157, no.
3796, p. 1554-1555, text-figs. 1-2.
Pessagno, E. A., Jr. and Miyano, K., 1968. Notes on the
wall structure of the Globlgerlnacea. Micropaleont
ology, vol. 14, no. 1, p. 38-50, pis. 1-7, text-figs.
1-2.
248
Reiss, Z. and Schneldermann, N., 1969. Ultranloro-
structure of Hoeglundlna. Micropaleontology, vol. 15»
no. 2, p. 135-i^» P-Ls. 1-3.
Silvestri, A., 1896. Foraminiferi pllocenlcl della
Provlncla dl Siena; Parte I. Accad. Pont. Nuovl
Llncei, Mem., vol. 12, p. 1-204, pis. 1-5.
Sorby, H. C., 1879. The anniversary address of the presi
dent. Proc. Geol. Soc. London, Session 1878-79;
Geol. Soc. London, Quart. Jour., vol. 35> appendix,
P. 56-93.
Sandberg, P. A. and Hay, W. W., 1967. Study of micro-
fosslls by means of the scanning electron microscope.
Jour. Paleo., vol. 41, no. 4, p. 999-1001, pis. 131-
132.
Towe, K. M., 1967. Wall structure and cementation In
Haplophragmoldes canarlensls. Cushman Pound. Poram.
Res., Contr., vol. 18, pt. 4, p. 147-151, pis. 12-13,
text-flg. 1.
Towe, K. M. and Clfelll, R., 1967. Wall ultrastructure In
the calcareous foraminifera: crystallographlc aspects
and a model for calcification. Jour. Paleo., vol.
41, no. 3, P. 742-762, pis. 87-99.
Williamson, W. C., 1858. On the Recent foraminifera of
Great Britain. London: Ray Soc. Pubis., p. 1-xx,
1-107, pis. 1-7.
Wood, A., 1949. The structure of the wall of the test
In the foraminifera; Its value In classification.
Geol. Soc. London, Quart. Jour., vol. 104, pt. 2,
p. 229-252, pis. 13-15.
Wood, A. and Haynes, J., 1957. Certain smaller British
Paleocene foraminifera, Part II— Clblcldes and Its
allies. Cushman Pound. Poram. Res., Contr., vol. 8,
pt. 2, p. 45-58, pis. 5-6.
APPENDICES
APPENDIX I
SAMPLE LOCALITIES
250
— V*
251
APPENDIX I
SAMPLE LOCALITIES
Allan Hancock Foundation Station Numbers
Station Depth
Humber Latitude Longitude (Meters)
Bollvlna argentea Cushman
4832 33° 45'
00" H 118° 27' 25"
W 357
6900
34° 03' 00" H 119°
14' 28" w
463
3168
33° 47'
40" H 118° 32' 10" w 576
11237 33° 38'
24" N 118° 22' 06" w 787
3170 33° 46* 38" N 118° 37'
02" w 842
6844
32° 37’ 00" N 118° 22' 18" w 1050
7193 34° 19' 55" N 120°
05'
30" w 532
Bollvlna snlssa Cushman
6849 32° 30' 58" H 117°
18' 34" w 335
3168 33° 47' 40" H 118° 32' 10" V 576
11237 33° 38'
24"
H 118° 22' 06" V 787
6638 32° 48' 10" H 118° 17' 50" V 988
6844
32o 37 •
00" H 118° 22' 18" V 1050
6842 32° 22' 50" H 117°
22' 12" w 1240
Bollvlna acuminata Natland
4844 34° 09' 15" H 119° 13'
50" w 12
Bollvlna oseudobevrlohl Cushman
6880 34° 15' 00" H 120° 15'
00" w 492
Bollvlna semlnuda Cushman
11773 24° 25' 03" H 110° 08' 16"
w 732
Islandlella lomltensls (Galloway and Vlssler)
6806 33° 56' 06M N 118° 52' 17" W 205 •*»
252
Station Depth
Number Latitude Longitude (Meters)
Islandlella tortuosa (Cushman and Hughes)
11238 33° AO' 20" N 118° 16’ 12" If 46
Florllue baelanlnatus (Cushman and Moyer)
4839 34° 10* 50" N 119° 17' 50" If 18
4813 34° 32' 20" N 120° 39' 00" If 51
4851 34° 05' 30" H 119° 05' 55" W 165
10066 35° 55' 12" N 119° 28' 50" W 274
Honlonella nlocenloa Stella Cushman and Moyer
4839 340 io' 50" S 119° 17' 50" W 18
4851 34° 05' 30" N 119° 05' 55" W 165
10066 35° 55' 12" N 119° 28' 50" W 274
5532 34° 05' 25" N 119° 14' 10" V 366
Pullenla salleburyl R. 6. and K. C. Stewart
5170 34° 20' 35" N 119° 45' 30" W 81
5572 34° 19' 42" N 119° 43' 00" W 83
Ghllostome11a ovoldea Beues
11234 33° 39' 48" N 118° 20' 38" If 632
R/V Anton Bruun Orulee 17 Station numbers
Bollvlna Interlunota Cushman
659D 12° 30' S 77° 28' W 420
Oancrle manriaun Plnlay
658D 12° 18' S 77° 29' w 210
658K 12° 24' 3 77° 27' W 287-300
6600 12° 43' 3 77° 27' W 735
Islandlella oallfomloa (Ouehman and Hughee)
66OC 12° 43' 3 77° 27' W 735
Station
Bomber Latitude Lpa&Uude
Oaeeidulina laevigata d'Orbigny
6590 12° 30' S 77° 28' ¥
Oaeeidulina pulohella d'Orbigny
6590 12° 30' 8 77° 28' W
Pullenla bulloldee (d'Orbigny)
665B 15° 36' 3 77° 34' ¥
Pullenla malklnae Coryell and Moeeoan
6590 12° 30' 8 77° 28' ¥
Hoeglnndlna eleaane (d'Orbigny)
6590 12° 30’ 8 77° 28' ¥
6600 12° 43' 8 77° 27' ¥
683J 33° 01' 8 72° 01' ¥
665B 15° 36’ 8 77° 34' ¥
666C 16° 43' 8 77° 14' ¥
ooncentrlca (Parker and Jonee)
6807 33° 38' 8 78° 48' ¥
U.8.H.S. Bltanln Station numbers
Lentloullna oultrata (Nontfort)
Bit. 92 32° 49' 8 71° 51' *
Maralnullna obeea Terquem
Bit. 126 57° 12' 8 62° 45' ¥
Oaeeidulina braelllenele Ouehman
Bit. 126 57° 12' 8 62° 45* ¥
Qyroldlna neoeoldanll Brotien
Bit. 126 57° 12' 8 62° 45* W
253
Oeptb
(Metere)
420
420
2640-2780
420
420
735
1130
2640-2780
2980
100
866
3800
3800
3800
254
Station Depth
Humber Latitude Longitude (Meters)
Bay of Blecay Samples
Bollvlna aenarlenels (Costa)
p-16 43° 37' 00" N 1° 50' 00” W 830
Cas8ldullna carlnata Sllvestrl
P-16 43° 37' 00M H 1° 50' 00" W 830
APPENDIX II
PLATB NUMBBR8
255
256
APPENDIX II
PLATE NUMBERS
All plates taken In the Electron Microscope Labora
tory of the Allan Hancock Foundation, University of
Southern California, are numbered. Plate numbers1 for
figures used In this paper are listed below.
This AHP Plate This AHF Plate
Paner Number Paper Number
2(1) 4207A 17(1) 4627C
2 2 4207B 17(2) 4618B
3(1)
4217 A
18(1)
4756A
3(2) 4217B 18(2) 4756B
4(1) 4575B 19(1)
4268-1
4(2) 4572B 19(2) 4267-1
5(1) 417 5A 20(1) 4857A
5(2) 4175B 20(2) 4856B
6(1) 3299-5 21(1) 4288-4
6(2)
3341B 21(2) 4293-2
7(1) 4222A 22(1) 4292-1
7 2 4222B 22(2 4301A
8 1) 4221A 23(1)
4302-2
8(2)
4221B 23 2) 4843A
9(1) 4158C
24(l) 486 5A
9(2) 4158B 24(2) 4865B
10 1
4159 A 25(1) 4970C
10(2) 4159B 25 2 4985A
11(1)
3292B
26(1)
4972B
11(2) 3293A 26(2) 5003A
12(1) 3310A
27(1) 5022C
12(2) 418 5B 27 2 5049A
i3 1
4250B 28(1) 5022B
13(2) 4303B 28(2 5051A
14(1) 4304A 29(1) 5063A
14(2) 4304b 29(2) 5026B
15 1) 4291A 30 5046A
15(2)
4293 B 32(1) 4357A
4290A 32 2 4318C
16(2) 4290B 33(1) 4318A
This AHP Plate
Paper Number
33(2) 4318B
34 1 4873A
34(2)
48740
35(1) 4384C
35 2 4383B
36(1) 4871C
36(2) 4871A
37 1 5005A
37(2) 49510
39 1) 5027C
39(2) 5050B
41 1) 4970B
41(2) 5004A
42(1) 5024C
42(2) 5049C
44 1) 4945A
44(2 4978A
45(1) 4586B
45(2) 4887B
46(1) 4584B
46(2) 45700
47(1) 4494A
47(2) 4886A
48(1) 4S8SA
4877B 48(2)
49(1) 4569B
49(2) 44930
50(1) 4845A
50(2) 48760
51(1)
5028A
51(2) 50280
52(l) 5045B
52(2) 50450
54(1) 49710
54(2) 4985C
55(1) 4945B
55(2) 4950A
56(1) 5001A
56(2) 5004C
58(1) 5025B
58(2) 5046C
59(1) 5027A
59(2) 5047B
257
This AHP Plate
Paper Number
60(1) 5023C
60 2 5050A
62(i ) 5002A
62(2) 4984B
63(1) 5002C
63(2) 5002B
63(3 4977B
65 1) 5018A
65(2) 5052B
66(3.) 4391C
66(2) 4391A
67 1 4419A
67(2) 44200
68(1) 4422C
68(2) 4426B
69(1) 4879A
69(2) 5064A
70 4429B

Linked assets

University of Southern California Dissertations and Theses

Conceptually similar

PDF

Neogene to recent Naticidae (Mollusca : Gastropoda) of the eastern Pacific

PDF

Oceanography And Late-Quaternary Planktonic Foraminifera, Southwestern Indian Ocean

PDF

The Stratigraphy, Foraminifera, And Paleoecology Of The Montesano Formation, Grays Harbor County, Washington

PDF

Late-Neogene Paleomagnetic And Planktonic Zonation, Southeast Indian Ocean - Tasman Basin

PDF

The Paleontology And Stratigraphy Of The Ely Group In The Illipah Area Ofnevada

PDF

Paleoecology And Stratigraphy Of Pre-Kaibab Permian Rocks In The Ely Basin, Nevada And Utah

PDF

Sedimentary History Of The Early Pliocene In The Los Angeles Basin, California

PDF

Facies Variation And The Miocene-Pliocene Boundary In Southern California

PDF

Ecology And Paleoecology Of Hudson Bay Foraminifera

PDF

Continental Slopes Of The World

PDF

The Origin And Distribution Of Glauconite From The Sea Floor Off California And Baja California

PDF

Distribution Of Foraminifera And Radiolaria In Sediments Of The Scotia Sea Area, Antarctic Ocean

PDF

Sediments Of The Southern California Mainland Shelf

PDF

Continental Margin From San Francisco, California, To Cedros Island, Bajacalifornia

PDF

Heavy Minerals In Sediments Of Southern California

PDF

Early Diagenesis In Southern California Continental Borderland Sediments

PDF

Sedimentology And Pleistocene History Of Lake Tahoe, California - Nevada

PDF

Heat Flow And Other Geophysical Studies In The Southern California Borderland

PDF

Marine Geology Of The Baja California Continental Borderland, Mexico

PDF

The Aleutian-Kamchatka Trench Convergence: An Investigation Of Lithospheric Plate Interaction In The Light Of Modern Geotectonic Theory

Asset Metadata

Creator Stapleton, Richard Pierce (author)

Core Title Ultrastructure Of Tests Of Some Recent Benthic Hyaline Foraminifera

Degree Doctor of Philosophy

Degree Program Geological Sciences

Publisher University of Southern California (original), University of Southern California. Libraries (digital)

Tag OAI-PMH Harvest,paleontology

Language English

Contributor Digitized by ProQuest (provenance)

Advisor Bandy, Orville L. (committee chair), Bils, Robert F. (committee member), Easton, William H. (committee member)

Permanent Link (DOI) https://doi.org/10.25549/usctheses-c18-430931

Unique identifier UC11362133

Identifier 7025064.pdf (filename),usctheses-c18-430931 (legacy record id)

Legacy Identifier 7025064.pdf

Dmrecord 430931

Document Type Dissertation

Rights Stapleton, Richard Pierce

Type texts

Source University of Southern California (contributing entity), University of Southern California Dissertations and Theses (collection)

Access Conditions The author retains rights to his/her dissertation, thesis or other graduate work according to U.S. copyright law. Electronic access is being provided by the USC Libraries in agreement with the au...

Repository Name University of Southern California Digital Library

Repository Location USC Digital Library, University of Southern California, University Park Campus, Los Angeles, California 90089, USA