Journal of Foraminiferal Research, V. 10, no. 1, p.
1-30, January 1980
QUANTITATIVE PALEOBATHYMETRY AND PALEOECOLOGY OF
THE LATE PLIOCENE-EARLY PLEISTOCENE FORAMINIFERA
OF LE CASTELLA (CALABRIA, ITALY)
M. L. BREMER,',~
M. BRISK IN^
ABSTRACT
The paleobathymetry and paleoecology of the Plio-
cene/Pleistocene boundary stratotype section at Le
Castella (Calabria, Italy) has been derived from a
study of benthonic and planktonic foraminifera.
A Q-mode principal component analysis simplified
the benthonic census data to a few assemblages which
were subsequently compared to the present day ben-
thonic population described by Parker (1958) in east-
ern Mediterranean surface sediments. The comparison
indicates that the Le Castella benthonic foraminifera
are similar to those occurring between 130-700 m in
the Mediterranean today-a range occupied by the In-
termediate Water Mass.
Benthonic foraminiferal principal component I1
summarizes the greatest dimension of variation in the
Le Castella faunal census data. The taxa Cassidulina
laevigata, Uvigerina peregrina, Bulimina costata, B .
aculeata + marginata and Hyalinea balthica dominate
one end-member of principal component 11; Cibici-
doides floridanus dominates the other.
The relative abundance of these two assemblages
changes just above the Pliocene/Pleistocene boundary
INTRODUCTION
The marine section at Le Castella (Calabria, Italy)
is the generally accepted Pliocene-Pleistocene bound-
ary stratotype. For this reason, the paleobathymetry
1 Department of Geology and Geophysics, Woods Hole Ocean-
ographic Institution, Woods Hole, Massachusetts 02543.
2 Department of Geology, University of Cincinnati, Cincinnati,
Ohio 45221.
AND W. A. BERGGREN'
(marker-bed). In the Pliocene section the fauna is
characterized by frequent fluctuations between the
Cassidulina laevigata-Uvigerina peregrina assem-
blage and the Cibicidoides floridanus assemblage; in
the Pleistocene unit, the Cassidulina laevigata-Uvi-
gerina peregrina assemblage dominates. The only re-
ported recent occurrence of C. floridanus with abun-
dances similar to those of Le Castella is in the Gulf of
Mexico.
Relatively high percentages of warm-water plank-
tonic foraminifera are associated with the Cassidulina
laevigats-Uvigerina peregrina assemblage, whereas
relatively low percentages are associated with the Cib-
icidoides floridanus assemblage. The integrated plank-
tonic and benthonic foraminiferal evidence delineates
a record of two climatic regimes in the stratotype. A
series of frequent climatic fluctuations, alternating be-
tween a characteristically Mediterranean climate and
one comparable to the Gulf of Mexico today, char-
acterizes the late Pliocene. The Pleistocene regime is
typified by a relatively constant climatic regime com-
parable to that of the Recent.
and paleoclimate of the section have been the object
of scientific interest for many years. Benson (1975)
studied the ostracodes in an attempt to reconstruct
depths and bottom temperatures; Marchetti (1975)
studied the pollen spectra to identify the climatic re-
gime; Lamb and Beard (1972) grouped planktonic fo-
raminifera into warm and cold categories to infer sur-
face-water temperatures. Emiliani and others (1961)
and Emiliani (1971) studied the isotopic composition
1
BREMER AND OTHERS

40'

0-.

-15

- 30

30" -45
I J
4 0' 20" 30°
60
FIGURE 1
Location of the section at Le Castella and of the surface sediment
samples of Parker (1958). After Parker (1958, p. 222). For depth 75
and location of samples see Parker (1958, p. 223-224).
90

in the test of planktonic foraminifera to delineate sur- 105

face temperatures. These studies suggest that the sed-

iments of Le Castella record frequent environmental
changes with a tendency for surface temperatures in
the Pleistocene to be lower than those in the Pliocene.
Although abundant and diverse, the benthonic fo-
raminifera of Le Castella have never been thoroughly
studied. They have a unique potential for the recon-
struction of the environment at the sea floor. By study-
ing both the benthonic and planktonic foraminifera of
the Le Castella section, it is possible to reconstruct
the environment of the deposition as well as the ocean-
ographic and climatic regime of that time. Specifically,
the objectives of this study are the following:
1) to estimate the paleobathymetry of Le Castella,
and
2) to reconstruct and interpret environmental
changes of Le Castella.
The first 'objective is accomplished by examining the
bathymetric distribution of the Le Castella assem-
blages in the present-day eastern Mediterranean fau-
nas described by Parker (1958). Q-mode principal
components analysis is used to define the assemblages
and examine their distribution. Although time con-
suming, this approach is inherently more objective and
complete because more species are considered and
relative species abundance information is considered
in greater detail.
The second objective is achieved by studying both
FIGURE 2
Location of the samples within the section at Le Castella. Major
foraminiferal and calcareous nannoplankton datum events are
shown. From Haq and others (1977, p. 485).
the planktonic and the benthonic foraminifera; in this
way it is possible to evaluate the relationship of change
at the sea surface to change at the sea floor.
THE STUDY AREA
Many of the well-exposed outcrops of Italian Plio-
cene/Pleistocene are located in the Catanzaro-Nicas-
tro structural trough of central Calabria (Bayliss,
1975). This trough is considered to have formed in the
early to middle Miocene with downwarping continuing
at least until early Pliocene (Burton, 1965, 1971).
There are small, isolated areas in this trough in
which the basement is overlain by sedimentary rocks
of Triassic to early Eocene age (Burton, 1965, 1971).
For the most part, however, the youngest sedim'entary
rocks found within the trough are conglomerates,
sandstones, clays and limestones of middle to late
Miocene age. The sands, clays and marls deposited in
the early Pliocene contain rich planktonic and ben-
thonic foraminiferal faunas thought to represent a rel-
atively deep-water environment (Bayliss, 1975). De-

2
 PLIOCENE-PLEISTOCENE IN ITALY

position of clays, marls and thin sandy horizons 1.0 -

continued until early Pleistocene. Today, many of
0
these Pliocene/Pleistocene sections are several hundred Ii
.9 - -17
meters above sea level. -6
The 155 m composite section at Le Castella consists *mi3
18
of gray silty claystones and thin- sandy horizons (Fig. .8 -
2). The thin sandy layer which occurs approximately
45 m below the top of the section is thought to be the
.7 -
lithostratigraphic marker bed of the Pliocene/Pleisto-
cene boundary as defined by Gignoux (1910). The
marker bed and some of the other sandy horizons .6 -
within the section are thought to have a turbidity cur-
rent origin (Bayliss, 1975). 0

The sediments at Le Castella contain shallow-water .5 -

benthonic foraminifera such as Elphidium sp., Florilus

boueanus, Asterigerinita mamilla and Ammonia bec- fil
c, ,4-
carii which today do not live in water deeper than 200 P
m. Also present are deeper-water species such as 0

Uvigerina peregrina, Cibicidoides floridanus and An- .3 -

gulogerina angulosa. These live in water greater than
200 m in the present-day eastern Mediterranean and -
.2
they are the dominant forms in the Le Castella fauna. 0 %
None of the shallow-water taxa ever exceeds 3% in
relative abundance, and these shallow-water species .i -
probably are not reworked or displaced. Thus, with
the use of multivariate statistical techniques designed
0 :
to emphasize abundant species, it is possible to extract -
0'
1

ecological information and gain valuable insight into

the paleoenvironment of this marine boundary strato-
type section.
- .I -
* *
.-
e
2- 0 : 0

I I I I
BIOSTRATIGRAPHY AND .C

-.6 -.4 -.2 0 .2 .4 .6 .6

CHRONOSTRATIGRAPHY
The biostratigraphic framework for this investiga-
tion is that of Haq and others (1977). Working with
the same set of samples, they found the following da-
tum events recorded in the Le Castella section: the
sequential FAD (first appearance datum) of four
species of the coccolithophore genus Gephyrocapsa,
the LAD (last appearance datum) of Discoaster brou-
weri, the LAD of the planktonic foraminifer Globiger-
inoides obliquus and the FAD of Globorotalia trun-
catulinoides. The positions of these datum events in
the section are shown in Fig. 2.
If the Gephyrocapsa caribbeanica FAD (1.62 Ma)
and the Discoaster brouweri LAD (1.65 Ma) are cho-
sen as the two most reliable datum events (Haq, per-
sonal communication) an average sedimentation rate
of 9.7 cd10,OOO years is obtained for this section.
Assuming constant sedimentation the time range of
the section is estimated to be 1.55 Ma to 1.75 Ma.
PC m
FIGURE 3
Plot of Le Castella samples on the second and third principal com-
ponent axes generated by the first principal components analysis
to show the composition of the samples in terms of the principal
component assemblages. The coordinates are factor loadings. The
samples fall into two clusters. Samples 18, 17, 13, 11 and 6 have
factor loadings for principal component I1 which are higher than
those of the other samples. They are characterized by high abun-
dances of Brizalina attica and Bulimina exilis.
DESCRIPTION O F THE FAUNA
B ENTHONIC FORAMINIFERA
The samples in this study were collected by F. Bar-
bieri in 1971. They are the same as those studied by
Rio (1974) and Haq and others (1977). Sixty samples
taken at approximately 2- to 3-m intervals span the
150-111 section (Fig. 2).

3
BREMER AND OTHERS

TABLE 1
Factor scores for the first ten Q-mode principal components. They represent the proportional contributions of each of the species to each of
the principal components.

Principal component
Taxon I 2 3 4 5 6 7 8 9 IO
-__
Ammonia beccarii .007 .018 -0.25 .011 .031 .188 - .046 - .005 ,117 - .476
Asterigerinita mammila .001 - .001 .001 .001 .001 -.001 -.001 - .ooo .004 - .007
Bigeneria nodosaria .002 - .002 - .005 .ow -.001 .001 - .ooo .ooo - .003 - .003
Brizalina albatrossi .013 .00s -.015 - ,006 - .003 ,020 - .006 -.OW ,004 - .040
Brizalina attica .017 .055 ~ .053 .05 1 .114 .055 .I92 -.018 ,049 ,022
Brizalina catanensis ,041 .I92 -.I50 .716 - .627 .082 .028 .010 -.I29 .008
Brizalina difformis .008 ,047 .030 .097 .I22 .ow ,047 .069 .064 .036
Brizalina dilitata .028 .065 .059 .lo3 ,228 .I61 - ,087 .031 -. 174 ,429
Brizalina pseudoplicata .001 .000 ,001 .ooo .ooo - .005 .001 - .ooo .001 ,001
Brizalina alata .oo7 .031 .008 .016 -.001 .018 -.OM .003 -.033 .035
Bulimina aculeata + marginata .195 ,266 - .289 - ,232 .084 .207 .053 .039 - .784 -.180
Bulimina costata .199 .267 -.701 -.201 - ,034 .246 -.I50 - .002 .477 ,036
Bulimina exilis .005 .020 - .044 .020 ,078 .158 .934 -. 147 ,116 -.001
Bulimina gibba .oo3 .012 - .003 .061 -.011 .088 .011 .003 .006 ,013
Cibicides boueanus .OS5 ,005 -.012 .041 -.016 .010 .001 .021 .041 .050
Cassidulina crassa .006 ,020 .016 .012 .016 - .002 - .007 .001 -.015 .013
Cassidulina laevigata .335 .421 ,196 .396 ,503 .021 - ,082 - ,002 .IO5 -.010
Cassidulina subglobosa .I02 ,028 .070 .OS6 .046 .068 .001 .020 - ,017 -. 180
Cibicides lobatulus .037 ,054 .049 ,072 .111 .034 .004 .015 .061 .011
Cibicidoides Joridanus ,701 - .655 - ,056 .078 - ,027 .082 .013 .003 - .066 ,059
Eggerella bradyi .004 -.01I - ,022 .022 - ,003 ,003 .001 .007 .002 - .005
Elphidium crispum .001 ,003 .003 .000 .001 .002 .001 - ,000 .005 - .007
Epistominella evax .002 .001 - .008 -.012 - .005 - .040 .146 .983 .05 1 - .022
Epistomeinella rugosa
convexa .023 - .006 .032 - ,004 .001 .012 - .008 -.001 .023 - .048
Florilus boueanum .005 .014 -.018 -.001 .030 .030 .008 .002 - .046 .016
Gavelinopsis praegeri .021 .061 .027 - .001 .031 -.008 -.009 .002 .004 .052
Globobulimina affinis .034 .025 - .041 - .010 .022 - ,041 - .001 .003 .001 .059
Globobulimina
pseudospinescens .OlS .046 - ,033 - .019 .024 - .007 - ,001 -.001 -.042 ,039
Gyroidina altqormis .001 -.001 .022 ,001 .001 .002 - .ooo .000 .Ooo - .003
Gyroidina umbonata .046 .018 .033 .077 .072 ,040 ,004 .004 ,016 - .062
Gyroidinoides neosoldanii .070 - .006 .028 .ooo -.016 .073 -.021 - .002 .030 -.170
Hanzawaia rhodiensis .001 - .ooo .003 -.001 - .003 .000 .001 - .ooo .003 .006
Hoglundina elegans .004 .009 .012 .003 .011 .010 - .006 .002 - .006 .037
Hyalinea balthica .IO8 .217 .401 - ,330 -.369 .464 .007 .020 .030 .360
Amphicoryna scalaris ,009 .033 ,026 - .038 - ,034 - .066 ,018 - ,006 .014 ,103
Melonis barleanus .I21 - .047 ,028 .062 .065 .083 - .030 ,016 .076 .348
Miliolinella circularis .001 .006 .005 .010 .012 - .008 .005 .ooo .008 - .002
Neoconorbina terquemi ,005 .017 .002 .010 .028 .020 -.013 .003 - .006 .035
Oridorsalis tener ,028 - .000 - ,074 - .008 .006 - .009 - .008 - ,004 .023 ,009
Planorbulina mediterranensis .001 - .002 ,000 .ooo .ooo .003 - .001 ,000 .001 - ,003
Planulina arimenensis .I23 - ,011 .092 - .090 -.117 .048 .006 .023 .062 .012
Pseudoclavulina crustata .013 - .004 .008 -.021 - .028 .015 - .002 -.001 .027 -.018
Pullenia bulloides ,024 .026 - .028 - .003 .010 -.017 .005 - .004 .007 - .006
Pullenia quinqueloba .012 .002 .013 - ,009 - .ooo .019 - .001 ,002 - .029 - ,033
Pyrgo anomala .002 .003 .002 - .003 - .001 -.010 .002 - .001 - .005 -.014
Quinqueloculina lamarckiana .007 .015 ,015 - .003 .007 - .022 .001 - .001 -.014 -.017
Rotamorphina involuta .001 ,001 - .003 .001 -.001 .001 - .ooo - .Ooo .001 -.001
Sigmoilina distorta .008 .011 ,022 - ,008 - ,005 - .008 ,001 -.001 .010 .003
Sigmoilina tenuis .007 .011 .015 - .011 - .014 -.010 .ooo - ,003 .022 - .007
Sigmoilopsis schlumbergeri .055 .027 .065 -.014 - .020 ,071 - .026 .001 .060 .030
Siphonina reticulatu .018 -.023 -.004 .005 -.016 .025 - .004 ,009 .017 .055
Sphaeroidina bulloides .I69 .075 .143 - .062 - .045 .068 - .032 - .004 ,086 - .275
Spiroloculina canaliculata ,004 ,005 .013 -.011 - .009 ,017 .002 ,002 -.016 .036
Textularia calva ,034 .017 .076 -.042 - .042 .141 -.033 ,007 ,079 -.012

4
 PLIOCENE-PLEISTOCENE IN ITAT v

TABLE 1
Continued.

Principal component
Taxon 1 2 3 4 5 6 7 8 9 10

Textularia conica .001 .002 ,002 -.001 -.002 -.008 .022 .007 .002 -.OM
Textularia sagittula ,021 -.003 .028 -.003 -.OOO .049 -.012 .002 ,005 -.055
Trifarina angulosa .158 ,131 ,349 -.057 -.072 ,188 -.046 -.005 .117 -.476
Uvigerina auberiana .OS1 -.047 -.048 .036 .020 .072 -.044 -.006 .113 -.242
Uvigerina peregrina .427 .336 .064 -.215 -.214 -.708 .126 -.049 .007 .138
Valvulineria bradyana .015 ,029 .012 ,083 .148 ,073 -.030 .017 -.050 .203

At least 150 benthonic foraminifera were randomly
selected from the greater than 150-micron fraction
(using a numbered grid and a random number table),
were identified and counted. The taxonomy is de-
scribed in Appendix l and the Le Castella benthonic
foraminiferal census data are given in Appendix 2.
After eliminating all species of benthonic foraminif-
era with maximum abundances of less than 1 percent,
the samples were described in terms of 60 species.
Some of the more abundant species are Uvigerina per-
egrina, Cibicidoidesfloridanus, Cassidulina laevigata,
Bulimina costata, B . aculeata + marginata, Brizalina
attica and B. exilis.
In order to simplify the data, a Q-mode principal
components analysis was performed. A description of
the mathematics of the technique may be found in
Davis (1973) and Daultrey (1976). Examples of the use
of this and similar techniques in paleoecology are
found in Imbrie and Kipp (1971), Haq and Lohmann
(1976), and Lohmann (1978). The computer program
CABFAC (Klovan and Imbrie, 1971) was used to per-
form the calculations.
The principal components analysis is advantageous
for two reasons. Firstly, it results in a substantial sim-
plification of the faunal census data while retaining
most of the information about the relative species
abundances. Secondly, it makes possible an objective
comparison of the Le Castella fauna to other data sets,
in this case, the Recent fauna of the eastern Mediter-
ranean.
An initial principal components analysis trans-
formed and reduced the 60 benthonic species to ten
assemblages. Principal component assemblages I1 and
I11 grouped the basic data into two clusters, one of
which consists of samples 18, 17, 13, 1 1 and 6 (Fig. 3).
The Q-mode principal components analysis is de-
signed to describe the major dimensions of an ellip-
soidal array of points (in this case, the samples are the
points and their coordinates are the species abun-
dances). If the technique is applied to a data set con-
sisting of two ellipsoids or clusters of points, then nei-
ther set will be described as well as when considered
separately. This is the case in our initial principal com-
ponents analysis; samples 18, 17, 13, 1 1 and 6 are dis-
tinctly different from all of the other samples in that
they have very high relative abundances of Brizalina
attica and Bulimina exilis. For this reason, those sam-
ples were considered separately and eliminated from
subsequent calculations.
After a second principal component analysis of all
samples except 18, 17, 13, 1 1 and 6, the samples
grouped into one cluster when plotted against principal
components I1 and 111. The factor scores and loadings
produced by this analysis are shown in Tables 1 and
2. The factor scores for the principal components rep-
resent the proportional contribution of each species to
each of the assemblages. The factor loadings for the
principal component assemblages represent the rela-
tive abundance of each of the faunal assemblages with-
in each of the samples.
Inspection of the factor scores shows that principal
component assemblage I (the vector mean) is defined
by abundant and ubiquitous species such as Cibici-
doides floridanus, Uvigerina peregrina, Cassidulina
laevigata, Bulimina costata and Bulimina aculeata
+ rnarginata. Principal component assemblage I1 is
defined by two end-member assemblages: one is dom-
inated by Cibicidoides floridanus and the other is
dominated by Cassidulina laevigata, Uvigerina pere-
grina, Bulimina costata, B . aculeata + marginata,
and Hyalinea balthica. The Cassidulina laevigata-
Uvigerina peregrina assemblage is abundant from
sample 45'to the top of the section (Fig. 4;Appendix
2).
PLANKTONIC
FORAMINIFERA
Parker (1958) and Ryan (1972) constructed warm-
cold climatic curves for the eastern Mediterranean
based on planktonic foraminifera (Parker identified the
species associated with relatively warm and cool en-
vironments qualitatively and Ryan identified them

5
BREMER AND OTHERS

TABLE 2
Factor loadings for the first ten Q-mode principal components. They represent the abundance of each of the assemblage in each of the
samples. We have followed Kaiser’s Rule (Cooley and Lohnes, 1971, p. 104) and retained principal components with eigenvalues greater than
unity. Although a specific environmental significance was attributed only to the second principal component, all ten principal components
were used to calculate the second index of similarity for the paleobathymetric estimate. In this way we maximize the proportion of the
benthonic foraminiferal faunal census data used to estimate paleobathymetry.

Principal factor loadings

Sample Comm. 1 2 3 4 5 6 7 8 9 IO

60 ,9671 .4413 .3057 - .6080 - .2843 - .0892 .3095 -.1252 ,0077 .3249 .0574
59 .9356 373 1 - .0407 .3215 - .0980 -.I550 .0210 .0199 -.0016 .0130 .1526
58 .9650 .8512 .2976 .3142 -.1579 -.I201 ,0268 .0063 - ,0020 ,0636 .0944
57 ,9044 .9191 .1130 - .OW8 - .0669 .0453 - .0536 .0125 .0022 -.1512 -.I203
56 3691 S603 .3618 .3776 - ,2697 -.1981 .3353 .0221 .0193 -.I673 .1686
55 .9405 ,7909 .3003 -.0232 -.0601 .1215 - .0296 .0448 .0104 - .4501 -.0164
54 3957 .7635 ,2362 .2547 - ,0963 -. 1092 - .3722 .0539 - ,0320 .0496 -.1612
52 .9299 .8861 .2765 .1629 - .0640 ,0198 .1248 - .0456 .0003 .I368 - ,0294
51 .9840 ,4876 .4919 -.1004 s755 - .3938 -.0631 .0139 -.mol - .0673 .0202
50 .7605 .3998 ,2765 .1760 .2632 .4507 .2560 -.1122 .0396 -.2021 .3169
49 .9397 ,7273 .5 146 .2475 .0807 ,2318 -.0216 -.0410 .0028 .0111 ,1487
47 .9232 .6770 .3949 .2157 -.2178 -. 1798 - ,3352 .0798 -.0235 .0216 .2508
46 .9585 .9376 ,0952 -.0819 - .0991 .0360 -.0192 .0127 - ,0008 -.1924 -.I221
45 ,9008 .8265 -.3406 .I743 - .0020 - .0842 .2077 -.0183 ,0079 ,0432 -.1374
44 .9570 .8661 - ,0373 .2758 - .2008 - .2859 - .0399 .0543 -.0021 .0034 .0467
43 .9471 .9090 ,1570 .0842 .0873 .I619 .0483 - .0626 - .0075 .I275 -. 1808
42 ,9669 .9366 -.1318 .2126 .0602 ,0460 .0978 -.0079 .0045 ,0015 -.IO85
41 ,9249 .8467 - ,3832 .0333 .1949 .1409 .0075 - ,0034 - .0004 - .0175 .0417
40 ,9524 .9650 -.0251 ,0474 .lo54 .0395 -.0193 - .0002 .OW6 - .0627 - .0358
39 .9354 A687 .0288 .2054 .2267 ,2416 -.0016 - .0500 - .0063 ,1249 - .0993
38 .9562 .9272 -.I957 -.0451 .0181 -.0103 - .2093 .0180 -.0189 .0804 .0697
37 .9637 3969 .2031 .2439 .I054 .0270 -.0118 -.0180 - .W92 .0718 - .2024
36 .9276 3817 ,1786 .5227 -.1630 - .2287 .4300 - .0495 ,0183 .1239 - .0463
35 .9627 ,8801 -.2330 .2425 -.1371 -.1983 .lo67 .0274 - ,0004 - .0397 - ,0565
34 .9812 .9669 - .0872 .I017 .040 1 .0305 - .0255 - .OH9 - .0095 .lo17 -.I199
33 .9525 .8974 .0500 .I027 .0154 .0086 - .3538 .0399 - .0257 .0654 .0455
32 .9012 3932 -.1309 .0879 .0689 .0940 .0477 - .0070 .0047 - ,0837 -.2359
31 .9615 .9463 -.I246 .0697 -.0124 -.0251 -.1623 ,0326 -.0126 - ,0243 -.1290
30 ,9695 3864 .I557 - .2789 -.1523 .0291 - .0820 .0256 - .0011 - .2203 - .0425
29 .9371 .9190 -.1238 -.1449 - .0253 .0478 - .0537 - .W03 .0057 -.IO13 .2005
28 .9694 .7866 .3870 -.3158 -.0617 .0628 -.I682 - .0526 -.0201 ,2438 ,0515
27 ,9908 ,0457 .I979 -.1543 ,7166 - .6234 .0815 .0359 .0092 -.1237 ,0135
26 .9992 .0629 ,0050 - .0231 -.0214 - .0077 - ,0494 .1507 .9830 .0498 -.0155
25 .9084 .9276 .0324 -.1570 - .0034 .I022 ,0028 - .0264 .OOO8 .o002 -.IO52
24 ,9646 .9181 -.2122 - .2442 - .0675 -.0167 .0926 - .0345 .0060 .0462 ,0157
23 .9748 .8545 -.0512 - .2427 -. 1484 ,0261 .0745 ,0248 .0198 - .3904 - .0397
22 .9658 .9048 - ,3233 - .0722 .0086 - .0076 - .0907 ,0233 - ,0004 -.0735 .1520
21 .9485 3719 - .4201 - ,0247 .0561 .0102 ,0820 -.0139 ,0039 ,0229 -.0216
20 .9782 .8474 .I310 -.4119 ,1725 - .0186 .1414 -.0418 .0019 .0949 -.0311
19 .9652 .8676 .1610 - .4029 .0893 -.1140 .0304 - .0178 - ,0018 .0341 - .0302
16 .9693 3798 - ,3925 - .0937 .097 1 .0566 .0548 -.0136 - ,0014 ,0922 .0900
15 .9596 ,9071 p.2514 -.0739 .I639 .0790 .0450 -.0410 .0017 .1323 .1175
14 .9491 ,6996 .4194 - .4770 -.1654 ,0417 - .0647 - ,0433 -.0141 .1482 .0165
12 .9706 3243 - .4958 - .0860 ,0389 - ,0596 -.0115 ,0130 ,0017 .0129 I1802

10 .9838 .748 I -.6173 -.OS11 .0797 -.0127 ,1275 .0110 ,0061 ,0029 .1165
9 .9573 .6554 .2688 - 3373 -. 1927 .0720 .I815 .1259 -.0105 -.1272 - .0560
8 .9832 .0097 .03 11 - .0496 ,0299 ,0978 .1643 .9514 -.1477 .I231 .0018
7 .9664 3666 - .3603 -.1059 - .0586 -.I215 -.1934 .0399 -.0158 ,0339 ,1257
5 .7408 .4027 .3830 .1705 .4055 .4547 .0097 .0890 ,0238 ,1407 .1578
Variance 63.78 7.73 6.00 3.62 3.15 2.49 2.10 2.03 1.99 1.43
Cum. Variance 63.78 71.51 77.51 81.14 84.28 86.77 88.87 90.91 92.89 94.32

6
 PLIOCENE-PLEISTOCENE IN ITALY

% with a factor and vector analysis). Parker used Has-

tigerina aequilateralis, H . pelagica, Globigerinoides
60
59
ruber and G . sacculifer, to indicate relatively warm
58 temperatures. Ryan found that H . aequilateralis, Glo-
-57 bigerinoides ruber and Globorotalia truncatulinoides
56
55 are associated with relatively warmer temperatures in
!% the Mediterranean. With the exception of G . trunca-
52 tulinoides, we have used Ryan’s warm-water species.
51 Because of its evolutionary appearance near the base
50 of the Pleistocene, G . truncatulinoides is not mean-
49 ingful as a paleoecological indicator in a section as old
as Le Castella.
46 At least 200 planktonic foraminifera in each of the
45 Le Castella samples were examined and counted. The
44 combined relative abundance of the two “warm”
species was calculated. Values for this combined
42
41 warm species range from 0 to 37%. Temporal varia-
40 tions in warm planktonic foraminifera are plotted next
to those of benthonic foraminiferal principal compo-
nent I1 in Fig. 4. The C. laevigata-U. peregrina as-
39 semblage is associated with low abundances of warm-
-38 water planktonic foraminifera and the C. floridanus
37 lu assemblage is associated with high relative abun-
36
35
34
s!
3
dances of warm-water planktonic foraminifera.

=32 3 PALEOECOLOGY
31
30 PREVIOUS
STUDIES
29
Much attention has been focused on the paleoecol-
28
-27 ogy, biostratigraphy and paleomagnetic stratigraphy
26 of the Le Castella section because of its importance
25
24 as the PlioceneIPleistocene boundary stratotype. The
23 results of some of the relevant paleoecological studies
22 are summarized in the following.
21
20
19 and C13/C12
018/016
Emiliani and others (1961) analyzed the oxygen iso-
topic composition of the tests of planktonic foraminif-
16 era. The 47 samples studied spanned the entire sec-
15
14
tion. They concluded that a major shift in temperature
could be discerned in the section, with a trend from
12

IO t
9
B The percent “warm” planktonic foraminifera is the combined abun-
7 dance of Hastigerina aequilateralis and Globigerinoides ruber. The
values for principal component assemblage I1 are factor loadings.
I I I I
+ . . ’ . . - ““‘(a- Negative factor loadings for benthonic principal component assem-
5
-.6 -.4 -.2 0 .2 .4 blage I1 indicate the presence of the C . floridanus assemblage and
positive factor loadings represent the presence of the C. laevigata-
FACTOR LOAD” U . peregrinh assemblage. Note that the C . floridanus assemblage
FIGURE 4 is associated with high abundances of warm water planktonic fo-
Plot of benthonic foraminiferal principal component assemblage I1 raminifera and the C. laevigata-U. peregrina assemblage is asso-
(A) and percent “warm” planktonic foraminifera (B) down section. ciated with low abundances of warm water planktonic foraminifera.

7
BREMER AND OTHERS
warmer temperatures in the Pliocene to cooler tem-
peratures in the Pleistocene. Emiliani (1971) subse-
quently studied both the carbon and oxygen isotopic
composition of the tests of planktonic foraminifera at
Le Castella. In this study, 63 samples were taken from
a 27-meter interval spanning the marker-bed which
represents the Pliocene/Pleistocene boundary. He
found that although the isotopic ratios varied, the
boundary itself was not characterized by a strong gra-
dient.
Planktonic foraminifera
Lamb and Beard (1972) determined the percentages
of warm-water and cold-water planktonic foraminifera
throughout the section at Le Castella. Their results
showed no striking temperature change across the
marker-bed, but their data suggested that, on the av-
erage, the Pliocene part of the section represents an
environment which was relatively warmer than that of
the Pleistocene part of the section.
Benthonic foraminifera
Several investigators have made qualitative esti-
mates of depth and temperature based on benthonic
foraminifera. Emiliani and others (1961) gave a quali-
tative description of the benthonic foraminiferal fauna.
By selecting eight of the species and considering the
depth at which these species live today, they estimated
that the Le Castella sediments were deposited in ap-
proximately 500 m of water. They also noted that the
fauna varies markedly from sample to sample and in-
terpreted this to mean that the environment underwent
numerous variations. They also related the appearance
of Hyalinea balthica in the Pleistocene to decreasing
temperatures. Using the same method as Emiliani
(1961), Bandy and Wilcoxon (1970) estimated that the
benthonic foraminiferal faunas represented water
depths of 600-1,000 m, or perhaps more. Lamb and
Beard (1972) concurred with the depth estimate of
Bandy and Wilcoxon (1970). Iaccarino (1975, p. 455)
indicated that the benthonic foraminiferal faunas and
ratios of planktonic to benthonic foraminifera sug-
gested a “reasonably deep open sea sedimentary en-
vironment. ”
Ostracodes
Benson (1975) presented ostracode evidence that
the sill depths of the Straits of Gibraltar and the Straits
of Sicily were deeper than today at the time the Le Cas-
tella sediments were deposited. In a study of many
Neogene Mediterranean sequences, Benson found
8
that psychrospheric ostracode faunas were present in
the Pliocene, gradually decreased, and disappeared in
the Pleistocene; and furthermore, that this faunal trend
could be seen at Le Castella. Benson postulated that
early Pleistocene sill depths at the Straits of Gibraltar
and the Straits of Messina were approximatly 1,500 m.
These sills are at approximately 300 m today. The os-
tracode evidence suggested that the change from 1,500
m to 300 m was gradual.
Pollen
Marchetti (1975, p. 472) studied the pollen spectra
of five samples (taken from an interval of 15 m which
spanned the marker-bed) from Le Castella. A se-
quence of climatic phases was delineated, beginning
with a “moderately cool period, through a dry-warm
one, to oceanic and oceanic-temperate conditions.”
PALEOBATHY
METRY
The paleobathymetry of the section can be estimat-
ed by examining the bathymetric distribution of the
Le Castella assemblages in the present-day eastern
Mediterranean. There are two major assumptions in
such an estimate.
The first assumption is that the ecologic response of
the benthonic foraminifera under study has not
changed significantly since the time of the latest
Pliocene. The second assumption is that at the time
the Le Castella sediments were deposited, the hy-
drography of the eastern Mediterranean was similar
to what it is today. This assumption is necessary be-
cause benthonic foraminifera respond not only to
depth but also to a variety of physical and chemical
factors that vary as a function of depth. In short, we
are assuming that the bathymetric distribution of the
Le Castella assemblages has not changed.
We have calculated two indices of similarity in our
comparison of the benthonic foraminifera of Le Cas-
tella with those of the present-day eastern Mediter-
ranean. The first index is simply the percentage of
benthonic foraminifera in one of Parker’s samples
which remains after eliminating all taxa which do not
occur at Le Castella. The second index is derived from
an estimate of the proportion of the fauna (in ‘one of
Parker’s samples after all taxa which do not occur at
Le Castella were eliminated) which can be attributed
to the Le Castella assemblages.
Values for the first index of similarity are given in
Appendix 3; they range from 0-100%. For surface
samples obtained between depths of 106 m and 1,884
m, an average of 77.4% of the benthonic foraminifera
 PLIOCENE-PLEISTOCENE IN ITALY

TABLE 3
An estimate of the factor loadings for the Le Castella samples in Parker’s (1958) present day eastern Mediterranean benthonic foraminiferal
faunas.

Sample Comm. 1 2 3 4 5 6 7 8 9 IO

BS32 .012 ,024 .038 .025 .046 .075 .025 ,004 .009 .035 ,004
4662A .029 .os5 .028 .036 .049 .080 .068 - .009 .013 .os5 ,086
4666 .465 ,268 .289 .lo4 .460 .I71 .125 - .OS9 ,016 .077 ,180
466 1 .005 ,027 .021 .028 .022 .033 .028 - .00S .004 .027 - ,007
4658 .323 ,245 .261 .068 .373 .118 .lo2 -.041 .014 .085 .131
4667 .476 .245 ,309 .006 .531 -.012 .128 - .024 ,016 .021 .145
4657A .402 .321 .334 .128 .340 .165 - .026 - .016 .005 .OS5 .139
4649A .490 .433 .429 .001 .205 -.I71 - .084 .046 .006 -.159 .113
4654A .016 .043 .047 ,050 .039 .066 .026 .002 .012 .055 .002
4668 .033 .075 .047 ,064 .OS6 ,084 .071 - .023 .013 ,048 .os 1
4654B .066 .065 .084 .044 .065 ,106 .lo5 .I30 .004 ,096 .020
4721A .670 .492 .459 ,249 .211 .310 - ,018 - .035 .001 .073 ,087
4652 .48 1 .497 .404 .159 - .003 - .065 -.174 .029 - .005 ,046 .093
BS21A .735 .551 ,446 .061 -.166 -.I73 -.269 .OS8 -.011 - ,061 .301
BS20 .762 .553 ,450 - .044 .006 -.271 -.316 .073 -.015 - .OS9 ,303
4657B ,742 ,544 ,442 -.loo -.164 - .295 -.217 .035 -.015 . I17 ,255
BS29 .732 .673 .373 - .306 -. 105 - ,053 .035 .036 .004 ,032 .168
4648 .875 ,627 ,425 - .296 - .267 -.217 - .046 - ,023 - ,006 .064 .256
4647 .523 ,486 .435 - .OS9 -.185 - .044 -. 180 ,026 - .006 - .095 .125
4657 .830 .576 .432 -.176 -.I18 - .244 -.421 ,041 - .029 .I12 .120
4718 .829 .510 .391 - .OS7 - .252 -.I83 - .553 .090 - .037 - .008 ,032
4694 .774 ,577 .422 - .402 - .242 -.191 .021 -.021 - ,004 .072 - ,007

4723 ,752 .550 .376 .118 -.154 -.I03 - .474 .OS6 - .028 ,071 .162
4687 ,886 .549 .359 - .03 1 -.286 -.196 -.531 .121 - .029 -. 186 .049
4685 .952 .695 .275 -.136 -.316 - .209 - .422 .loo - .024 -.171 .114
4664 .648 .597 .218 -.251 -.153 - .028 - .os5 - .OS9 - .002 .I80 .343
4660 309 .628 .278 - ,200 -.221 -.128 - .298 - .002 -.021 .I77 ,335
47 14 ,138 ,276 ,068 .022 -.033 .013 .039 .OS6 ,006 ,007 .226
4656 .757 .436 .308 ,051 -.186 -.172 -.599 .I02 - ,038 .022 ,182
4713 .064 ,233 .OS8 - .002 - ,039 -.012 .020 - ,022 .005 .046 - ,034
BS9 .459 .422 .256 .097 -.160 -.I54 -.385 .060 - .029 ,057 - ,027

4654 .192 .235 .150 .OS3 - .OS7 -.035 - .255 .03 1 - ,012 - .002 .202
4686 .907 .495 ,340 .032 -.241 - .205 - ,641 ,125 - .040 - .073 ,107
4702 .lo8 .173 - .020 ,039 .038 .034 ,067 -.031 .001 .090 ,245
4650 .148 .206 .060 .038 .011 ,009 - ,001 - .028 ,007 .IO5 ,297
4670 .246 .364 .220 .020 -.I15 - .064 - ,211 .034 --,012 -.035 ,004
467 1 .528 .478 .173 .073 -. 138 -.I51 - ,441 .074 - .026 .003 .145
4693 .057 .146 ,057 .036 - ,040 -.os1 - ,042 - .001 -.010 .033 -.155
4699 .030 ,150 - .006 - .008 .003 ,004 .027 -.017 ,002 .062 .056
4688 .043 .071 .oos ,025 - ,001 -.016 .074 - ,022 - ,002 .030 -. 172
4716 .032 .089 ,010 .025 - ,004 - .014 .048 - ,019 .003 .025 -.141
4705 .040 .069 - ,006 .028 ,000 -.015 .071 -.021 - .002 .029 -.166
4673 .034 ,082 - .009 .027 ,005 - .009 .079 - .024 - .ooo .036 -.137
4678 ,011 .034 .002 - ,004 - .005 - ,007 .040 -.012 -.001 .012 - ,088
4683 .004 .017 .004 -.013 - .006 - .004 .024 - .007 -.001 .005 - .049
4689 .371 ,434 .077 .032 -.118 -.143 - .369 .066 - .030 .007 - .004

4698 .015 .064 .048 - .OS9 - .036 - .022 - .009 - .009 - .004 .041 - .039
470 1 ,002 .013 .005 -.015 - ,006 - .003 .020 - ,006 - ,001 .004 - .040
468 I .002 .013 .005 -.015 - .006 - ,003 .020 - .006 p.001 .004 - .04O

4710 .012 .062 .047 - .039 - ,034 - .023 - .033 - .002 - .005 .027 - .028
471 1 .001 .008 .011 .022 - ,007 - .004 - .009 .002 -.001 ,010 .003
4708 .002 .013 .005 -.015 - .006 - ,003 .020 - .006 -.001 ,004 - .040

4676 .210 .096 -.012 .029 ,102 .151 .110 - .042 .024 .018 ,390
Variance 13.628 6.471 1.321 3.OS5 1.680 5.705 .251 ,026 .612 2.591
Cum. Variance 13.628 20.099 21.420 24.475 26.155 31.860 32.111 32.137 32.749 35.340

9
 BREMER AND OTHERS

COMMUNALITY recorded by Parker also was included at Le Castella.

For water shallower than 106 m, the average is 53.7%;
0 .2 .4 .6 .8 1.0 for water deeper than 1,884 m, the average is 25.4%.
0 The second index of similarity is the communality
obtained in a principal components comparison (Klo-
van and Imbrie, 1971, p. 64) of the benthonic forami-
100 nifera of Le Castella and that of the present-day east-
ern Mediterranean. The communality is an estimate of
the proportion of the fauna which can be attributed to
200 the Le Castella assemblages. An index of 1 would in-
dicate that all the fauna in one of Parker’s samples
could be attributed to the Le Castella assemblages; an
300 index of 0 would indicate that none of the fauna in one
of Parker’s samples could be attributed to the Le Cas-
0
tella assemblages. Figure 5 is a plot of Parker’s sam-
400 ples against depth and the second index of similarity.
The similarities rise sharply at about 130 m, remain
high until they begin to decrease rapidly at about 700
500 m, at which depth their values remain consistently
0
4 low down to 3,974 m.
600 Both the first and second index of similarity are high
within the depth range of 130 to 700 m. That is, within
2fJ 700
this range the Le Castella and surface faunas have a
maximum number of species in common and the pro-
portions of these species are most similar. Thus, the
% 800
water depth during deposition of the Le Castella sed-
iments is inferred to have been between 130 and 700
2
\ m.
This depth estimate is significantly shallower than
3
L
900 that of Bandy and Wilcoxon (1970) and Lamb and
Beard (1972); it is consistent with that of Emiliani and
others (1961). Using an approach similar to that of Ban-
dy and Wilcoxon (1970), we were able to generate a
number of different depth estimates simply by select-
ing different taxa and emphasizing different depth dis-
tribution data sets. Figure 6 shows, for example, that
their technique can be used to produce a depth esti-
mate of -350-500 m which is consistent with ours
(-130-700 m). We think that our approach for esti-
mating bathymetry is inherently more objective be-
cause it uses a much greater number of taxa, and it
considers relative species abundance information in
greater detail.

1:
4O0OL
t t
larity indicates the proportion of the benthonic foraminifera in one
of Parker’s samples which can be attributed to the Le Castella
assemblages. The plot shows the bathymetric distribution of this
FIGURE 5 index of similarity. The dashed lines indicate the approximate depth
The second index of similarity (communality) versus depth for Par- region in which the communalities are high (Le., the Le Castella
ker’s (1958) surface sediment samples. This second index of simi- and surface sediment samples are similar).

10
 PLIOCENE-PLEISTOCENE IN ITALY

TEMPORAL
VARIATION
OF THE ENVIRONMENT
Because semi-enclosed seas are extremely sensitive
to climatic change, it is important to examine the as-
sumption that the hydrographic structure of the Med-
iterranean was similar at the time of deposition of the
Le Castella sediments to what it is today. For in-
stance, Ryan (1972) showed that the Pleistocene sap-
ropels of the Mediterranean probably were deposited
during periods of stagnation which developed at times
of glacial retreat. The fresh meltwater at the surface
would produce intense stratification of the water col-
umn and stagnation of the lower layers.
If the environment of the eastern Mediterranean
changed significantly during the deposition of the Le
Castella sediments, the changes should presumably be
recorded in the benthonic foraminiferal faunas. Prin-
cipal component assemblage I1 represents the major
dimension of variation in the fauna. As stated previ-
ously, it summarizes the reciprocal variation between
two assemblages. The taxa in the C. laevigata-U. per-
egrina assemblage occur in the present-day eastern
Mediterranean with relative abundances similar to
FIGURE 6
those of Le Castella. The C. floridanus assemblage is
The present bathymetric range of selected Le Castella taxa. Some
much more abundant at Le Castella than in the pres- of the rarer taxa are included. The dashed portion of the ranges
ent-day eastern Mediterranean. The greatest observed indicates depths at which the taxa become much less abundant.
relative abundance of C. Poridanus at Le Castella is The stippled area is the region where the depth ranges of all the
53% whereas in the present-day eastern Mediterra- taxa occur with relative abundances similar to those of Le Castella.
nean it is only 10%. Where do benthonic foraminiferal The sources of the depth ranges of each of the species are the
following: Martinotiella communis, Murray (1970), M u m (1976);
faunas with relative abundances of C. floridanus Sigmoilopsis schlumbergeri, Parker (1958), Lamy (1973); Uvigeri-
greater than 30% occur today? The only such record nu auberiana, Mum and others (1976); Sphaeroidina bulloides,
is that of Bandy (1956) (Cibicides pseudoungerianus Berggren, 1975, Berggren and others (1976), M u m (1976); Pullenia
sensu Bandy = Cibicidoides floridanus of this paper). bulloides, Berggren and Haq (1976), Berggren and others (1976),
He reported C. floridanus in the northeastern Gulf of M u m and others (1976); Melonis soldanii, Parker (1958); Melonis
barleeanus, Parker (1958); Planulina arimenensis, Mum and others
Mexico with relative abundances up to 47% (the max- (1976); Bulimina marginata, Parker (1958), Colom (1974), M u m
imum at Le Castella is 53%). and others (1976);Bulimina aculeata, M u m and others (1976); Bri-
The factor loadings for principal component assem- zalina albatrossi, Parker (1954), Parker (1958); CibicidoidesJlori-
blage I1 are plotted in Fig. 4. The appearance of this danus Parker (1958); Uvigerina peregrina, Phleger and Parker
curve changes at sample 46, 7.2 m above the marker- (1951), Mum and others (1976); Bulimina tenuata, M u m and others
( 1976).
bed. Above sample 46, the C. laevigata-U. peregrina
assemblage dominates (factor loadings in this portion
of the curve range from -.0407 to 3146). Below sam-
ple 46, the curve is characterized by rapid fluctuations other does not coincide with, but is very close to, the
between C. laevigata-U. peregrina and the C . flori- Pliocene/Pleistocene boundary (above the marker-
danus assemblage (factor loadings in this portion of bed).
the curve range from -.6173 to ,4194). This variation The high relative abundances of Cibicidoides flori-
in principal component I1 suggests that significant danus in the lower part of the section suggest that at
changes occurred in the paleoenvironment of the east- times during the Pliocene, conditions may have exist-
ern Mediterranean during the deposition of the Le ed at Le Castella which are not found in the eastern
Castella sediments. If these variations are attributed Mediterranean today.
to local climatic change, then there seem to have been Samples 18, 17, 13, 11 and 6 with high abundances
two climatic regimes in the eastern Mediterranean dur- of Brizalina attica and Bulimina exilis (which were
ing Le Castella time. The shift from one regime to the deleted from the final principal components analysis)

11
BREMER AND OTHERS
also suggest that the environment of the eastern Med-
iterranean may have been different at times during the
Pliocene. These two taxa do tlot occur in the present-
day eastern Mediterranean with abundances like those
of the Le Castella. In fact, no Recent analog has been
found.
In our estimate of the paleobathymetry of Le Cas-
tella, we assumed that at the time the Le Castella sed-
iments were deposited, the hydrography of the eastern
Mediterranean was similar to what it is today. At the
times that the C. laevigata-U. peregrina assemblage
dominated the benthonic foraminiferal fauna, this as-
sumption seems reasonable. When the C. floridanus
assemblage dominated the fauna, the environment at
Le Castella probably was different from any in the
present-day eastern Mediterranean. Bandy (1956) re-
corded C. floridanus in relative abundances greater
than 25% in depths ranging from 200 m to 600 m; this
range is similar to our estimated depth range (130-700
m) for Le Castella. For this reason, although the hy-
drography at Le Castella probably did change, we
think that our depth estimate for Le Castella is rea-
sonable.
CLIMATIC
VARIATION
By comparing the curves for benthonic foraminiferal
principal component I1 and warm-water planktonic fo-
raminifera, it is possible to study the relationship be-
tween changes at the sea surface and changes at the
bottom (Fig. 4). Results show that the temporal vari-
ations of warm planktonic foraminifera parallel those
of the C. laevigata-U. peregrina and C . floridanus
assemblages. The C . laevigata-U. peregrina assem-
blage is associated with the low percentages of warm-
water planktonic foraminifera and the C. floridanus
fauna is associated with high percentages of warm-
water planktonic foraminifera (Fig. 4).
Ciaranfi and Cita (1973) used planktonic foraminif-
era to delineate climatic change in the Pliocene of the
Mediterranean. Two of the warmest peaks on their
inferred climatic curve occur in the latest Pliocene.
Separating the two warmest peaks is a relatively cool
interval of short duration. These warm peaks may cor-
relate with those of Le Castella; both occur just below
the Globorotalia truncatulinoides FAD (Fig. 2). The
frequent climatic changes at Le Castella coupled with
the fact that the sedimentation rate, probably, was not
constant, make peak-to-peak correlation with the cli-
matic curves of Ciaranfi and Cita (1973) and others
very difficult.
The integrated benthonic and planktonic foraminif-
eral data appear to delineate two climatic regimes dur-
12
ing the deposition of the Le Castella sediments. The
demarcation point between these two regimes is very
close to the Pliocene/Pleistocene boundary. The Plio-
cene regime was characterized by frequent climatic
changes, while the Pleistocene regime was compara-
tively stable.
The planktonic foraminiferal faunas in the Pleisto-
cene unit indicate relatively cool temperatures. The
similarity of the associated benthonic foraminifera
with those at intermediate depths in the present-day
eastern Mediterranean suggests that the Pleistocene
environment at Le Castella was somewhat similar to
the Recent.
In contrast, the planktonic foraminifera of the Plio-
cene unit indicate frequent fluctuations between a rel-
atively cooler and a relatively warmer climate. The
benthonic foraminifera suggest that these climatic
fluctuations may have had a direct effect on the hy-
drography at depth.
The cooler phases in the Pliocene were associated
with a benthonic foraminiferal fauna similar to that of
the Le Castella Pleistocene and the Recent eastern
Mediterranean at intermediate water depths. The
warmer phases were associated with a rich benthonic
foraminiferal fauna dominated by C. $oridanus. Fau-
nas such as these have not been found in the present-
day eastern Mediterranean. Today similar faunas are
found only in the northeastern Gulf of Mexico (Bandy,
1956). Therefore, during the warmer phases of the
Pliocene, the Le Castella environment in the eastern
Mediterranean may have been more like that of the
Gulf of Mexico. To characterize the environment of
this time, it is instructive to examine the differences
between the Gulf of Mexico (following the classifica-
tion of the adjacent seas of the North Atlantic of Sver-
drup and others [1942] we will include the Caribbean
as part of the Gulf of Mexico) and the Mediterranean.
They can be summarized as follows:
1. They are at different latitudes.
2. They differ in their degree of isolation from the
Atlantic Ocean. The minimum sill depth between
the Mediterranean and the Atlantic is approxi-
mately 300 m (Sverdrup and others, 1942, p. 637-
646). The sill depth between the Gulf of Mexico
and the Atlantic is approximately 1,500 m. How-
ever, Benson (1975) gave ostracode evidence that
indicates that early Pleistocene sill depths at the
Straits of Gibraltar and the Straits of Messina
were approximately 1,500 m.
3 . Only Atlantic surface water flows into the Med-
iterranean (Lacombe and Tchernia, 1972, p. 32),
whereas Atlantic surface water and a consider-

able amount of Subantarctic Intermediate Water
flow into the Gulf of Mexico.
4. The structure of the water column between 200-
800 m is different in the Gulf of Mexico and the
Mediterranean. At these depths, temperatures in
the Gulf of Mexico range from 6-18"C, and in
the Mediterranean they range from 13"-15"C.
Salinities in the Gulf of Mexico range from 34.90-
36.70%0,and in 'the Mediterranean they range
from 38.45-38.94%0.
Temperature and salinity ranges in the Mediterra-
nean are narrower because vertical convection occurs
more frequently and extends to greater depths than it
does in the Gulf of Mexico (Sverdrup and others,
1942). The salinity differences are attributable to cli-
mate. Van Loon (1972, p. 109) gives a world map of
mean precipitation minus evaporation. Values of the
evaporation surplus in the Mediterranean range from
-50 to more than -100 cm per year, as compared to
-25 cm in the Gulf of Mexico. These differences are
related to large-scale features of atmospheric circula-
tion. The Mediterranean is within the narrow belt of
subtropical highs, the horse latitudes. The Gulf of
Mexico lies south of this belt (Trewartha, 1968, p. 88).
In the subtropical cell the high pressure pattern is
maintained by descending air. This air warms adi-
abatically and develops a low relative humidity (Donn,
1975, p. 241). This in turn leads to high rates of evap-
oration in the Mediterranean.
Any model invoked to explain the variation in the
Le Castella benthonic and planktonic foraminiferal
faunas must take into account the fact that the changes
observed are periodic. For this reason the gradual tec-
tonic uplift of the sills and exclusion of deep Atlantic
water postulated by Benson (1975) would not explain
the frequent oscillations in the foraminiferal faunas
(Fig. 4). At least two possible mechanisms would pro-
duce periodic changes: (1) eustatic fluctuations in sea
level and (2) climatic change.
A change in sea level today would have a pro-
nounced effect on the Mediterranean because it would
change the effective sill depth. The surface and inter-
mediate water masses are more sensitive to sea level
change when the sill is closer to the surface. If, as
Benson (1975) proposes, the sill depths during Le Cas-
tella times were greater, then the fluctuations of sea
level would have had much less effect than they would
today.
We suggest that the foraminiferal faunas varied as
a function of climate. The Mediterranean is extremely
sensitive to climatic change because it is a semi-en-
closed sea and it lies within the narrow belt of the
PLIOCENE-PLEISTOCENE IN ITALY
subtropical high. The benthonic foraminiferal assem-
blages of the Pliocene unit at Le Castella suggest that
the subtropical high was episodically displaced north-
ward. Climatic conditions at Le Castella, therefore,
periodically alternated from Mediterranean to tropical
as in the Gulf of Mexico.
MEDITERRANEAN INTERMEDIATE WATER
The depth estimate in this study corresponds rough-
ly to the average depth of 130-600 m given for the
Mediterranean Intermediate Water by Sverdrup and
others (1942). Hooper (1969) performed a factor anal-
ysis of Parker's samples of surface sediment with sim-
ilar results. He noted that the faunas of Parker's sam-
ples taken between 201 and 799 m were similar and
that this depth range corresponded roughly to the av-
erage depth of 200-700 m given by Pollak (1951) for
the Mediterranean Intermediate Water. But the bathy-
metric extent of the Mediterranean Intermediate
Water is quite variable.
Is there a consistent relationship through time be-
tween the benthonic foraminifera and the water mass
in which they lived? All attempts to relate the principal
component and factor assemblages and water mass
origin of Parker's samples have proven unsuccessful.
Assuming that the distribution of benthonic foraminif-
era is related to hydrography, there is a plausible phys-
ical explanation for this lack of success. Pollak (1951)
showed that the Adriatic is a source of Mediterranean
Intermediate Water. He found that the water structure
is directly affected by the prevailing weather regime
and that it changes both seasonally and yearly. In the
presence of such variable hydrography, it is probable
that a benthonic foraminiferal fauna from a sample
(which represents an average of the faunas of 100
years or more) could be correlated not with the hy-
drography at one point in time, but with the average
hydrography. We can speculate that the close similar-
ities between 130 and 700 m may represent an average
faunal response to the average vertical extent of the
Mediterranean Intermediate Water. It is, however,
difficult to test.
SUMMARY AND CONCLUSIONS
1. Using Q-mode principal components analysis, a
quantitative comparison is made between the ben-
thonic foraminiferal assemblages of Le Castella and
those of the present-day eastern Mediterranean de-
scribed by Parker (1958). This comparison indicates
that the Le Castella taxa were similar to those found
today between 130-700 m-a range occupied by the
Intermediate Water Mass.
13
BREMER AND OTHERS
2. No direct relationship between the benthonic fo-
raminiferal assemblages and water mass is observed.
We suggest that this is due to the variable hydrography
of the eastern Mediterranean at intermediate depth.
The paleobathymetric results correspond with the av-
erage depth of the Mediterranean Intermediate Water.
3. The Pliocene unit is characterized by frequent
alternations of the Cassidulina laevigata-Uvigerina
peregrina assemblage and Cibicidoides floridanus as-
semblage. The Pleistocene unit is characterized by the
dominance of the C. laevigata-U. peregrina assem-
blage .
4. In places Cibicidoides floridanus occurs in the
Pliocene unit of Le Castella in relative abundances
greater than those of the present-day eastern Mediter-
ranean. The only record of Cibicidoides Jloridanus
with abundances similar to those of Le Castella is in
the Gulf of Mexico. Bandy (1956) recorded C. pori-
danus in relative abundances greater than 25% in
depths ranging from 200-600 m in the northeastern
Gulf of Mexico; this is similar to our estimated depth
range (130-700 m) for Le Castella.
5 . The C . laevigata-U. peregrina assemblage is as-
sociated with low relative abundances of warm-water
planktonic foraminifera whereas the C. floridanus as-
semblage is associated with high relative abundances
of warm-water planktonic foraminifera.
6. The integrated planktonic and benthonic fora-
miniferal data delineate two climatic regimes. The de-
marcation point between these two regimes is situated
just above the marker-bed for the Pliocene/Pleistocene
boundary. The Pliocene regime was characterized by
frequent fluctuations between cooler and warmer sur-
face-water temperatures. During the cooler phases,
the hydrography was similar to that of the present-day
eastern Mediterranean. During the warmer phases, the
hydrography was similar to the present-day Gulf of
Mexico. The Pleistocene regime at Le Castella was
characterized by relatively cool surface-water temper-
atures and a hydrography similar to that of the present-
day eastern Mediterranean.
7. It is postulated that the Le Castella foraminiferal
faunas recorded episodic northward displacements of
the subtropical high during Pliocene time, resulting in
an alternation of climatic conditions.
ACKNOWLEDGMENTS
This paper was submitted by M. L. Bremer as par-
tial fulfillment of the requirements for a Master of
Science degree from the University of Cincinnati.
The suggestions and contributions of A. Chave, C.
Denham, J. Harrell, M. McCartney, R. Miller and L.
14
Tjalsma were very helpful. We would like to thank G.
P. Lohmann for his support, guidance and construc-
tive criticism, and B. U. Haq for his aid in establishing
the biochronology . Special thanks also are extended
to Frances Parker for her help in developing the taxo-
nomic framework, and to Ruth Todd and Wylie Poag
for their help with the taxonomy.
Particular thanks are due to Bob Commeau and Jack
McLane of the Office of Marine Geology, Atlantic and
Gulf of Mexico Branch, U.S.G.S., for their care and
skill in taking the electron micrographs. John Grover,
Anne Edwards and Harlow Farmer are thanked for
their help and encouragement. Many thanks are due
to Sandy Tonge of the Woods Hole Oceanographic
Institution and Jean Carroll of the University of Cin-
cinnati who typed the manuscript.
This research was supported by G.S.A. Penrose
Bequest Grant #2134-76, a Woods Hole Oceanograph-
ic Institution Summer Student Fellowship, the Grad-
uate Research Council Summer Fellowship from the
University of Cincinnati, and support from the Uni-
versity of Cincinnati General Fund; also contract 251
53 from a group of oil companies (Chevron, Exxon,
Marathon, Mobil and Shell) in support of Cenozoic
benthonic foraminiferal studies at Woods Hole Ocean-
ographic Institution. This is Woods Hole Oceano-
graphic Institution Contribution Number 4227.
REFERENCES
BANDY,0. L., 1953, Ecology and paleoecology of some California
foraminifera; Part I-The frequency distribution of Recent fo-
raminifera off California: Journal of Paleontology, v. 27, no. 2,
p. 161-183.
, 1956, Ecology of foraminifera in northeastern Gulf of Mex-
ico: U.S. Geological Survey Professional Paper 2746, p. 179-
203.
and WILCOXON, J. A., 1970, The Pliocene-Pleistocene
boundary in Italy and California:Geological Society of America
Bulletin, p. 2939-2948.
BARKER, R. W., 1960, Taxonomic notes on the species figured by
H. B. Brady in his report on the foraminifera dredged by H.
M. S. Challenger during the years 1873-1876: Society of Eco-
nomic Paleontologists and Mineralogists Special Publication 9,
238 p.
BATSCH, A. I. G. C., 1791, Testaceorum arenulae marinae tabulae
sex . . . (Sechs Kupfertafeln mit Conchylien des Seesandes).
Jena: University Press, p. I , 4, in Ellis, B. and Messina, A.,
ed., 1940, Catalogue of Foraminifera: American Museum of
Natural History (supplements post-1940).
BAYLISS, D. D., 1975, The stratigraphy of the type section for the
Pliocene/Pleistocene boundary in Italy, in Saito, T., and Burc-
kle, L. H . , eds., Late Neogene Epoch Boundaries: New York,
Micropaleontology Press, American Museum of Natural His-
tory, p. 31-40.
BELFORD, D. J . , 1966, Miocene and Pliocene smaller foraminifera
from Papua and New Guinea: Australian Bureau Mineralogical
Research, Geology Geophysics Bulletin, v. 79, 305 p.

BENSON, R. H., 1975, Ostracodes as indicators of threshold depth
in the Mediterranean during the Pliocene, in Stanley, D. J.,
ed., The Mediterranean Sea, A Natural Sedimentation Labo-
ratory: Stroudsburg, Pennsylvania, Dowden, Hutchinson and
Ross, Inc., p. 63-75.
BERGGREN, W. A., BENSON,R. H., HAQ, B. U., RIEDEL,W. R.,
SANFILIPPO,A., SCHRADER, H. J., and TJALSMA,R. C.,
1976, The El Cuervo Section (Andalusia, Spain): Micropaleon-
tologic anatomy of an early Late Miocene lower bathyal de-
posit: Marine Micropaleontology, v. l , no. 2, p. 195-247.
, and HAQ, B. U., 1976, The Andalusian Stage (Late Mio-
cene): biostratigraphy, biochronology and paleoecology: Pa-
laeogeography, Palaeoclimatology, Palaeoecology, v. 20, p. 67-
129.
BRADY,H. B., 1884, Report on the foraminifera dredged by H. M.
S. Challenger, during the years 1873-1876: Reports of the Sci-
entific Results of the Voyage of H.M.S. Challenger, v. 9 (Zo-
ology), 814 p.
BROTZEN, F., 1936, Foraminiferen aus dem Swedischen untersten
Senon von Ericksdal in Schonen: Sveriges Geologiska Under-
sokuing, ser. C. no. 396, p. 5-206.
BURTON, A. N., 1965, Geological mapping of the province of Cal-
abria, southern Italy at a scale of 1:25000: Geological Society
London, Proc., no. 1619, p. 1-5.
, 1971, Note on the alpine orogeny in Calabria, southern
Italy: Geological Society London, Quarterly Journal, v. 126, p.
369-381.
CIARANFI, N., and CITA,M. B., 1973, Paleontological evidence of
changes in the Pliocene climates, in Ryan, W. B. F., Hsu, K.
J., and others, 1973. Initial Reports of the Deep Sea Drilling
Project, Volume XIII, Washington ( U S . Government Printing
Office), p. 1,387-1,400.
COLOM,G., 1974, Foraminiferos IbBricos: Investigaci6n Pesquiera,
Barcelona, 245 p.
CUSHMAN, J. A., 1918, The foraminifera of the Bryam calcareous
marl at Bryam, Mississippi: U.S. Geological Survey Profes-
sional Paper 129-E, p. 87-105.
, 1923, The foraminifera of the Atlantic Ocean. Part 4, La-
genidae: United States National Museum Bulletin, pt. 4, p. 1-
228.
, 1931, The foraminifera of the Atlantic Ocean, Part 8, Ro-
taliidae, etc.: U.S. National Museum Bulletin 104, pt. 8, p. 1-
179.
, 1937, A monograph of the subfamily Virgulininae of the
foraminiferal family Buliminidae: Cushman Laboratory for Fo-
raminiferal Research Special Publication 9, p. 1-228.
, 1939, A study of the foraminifera contained in cores from
the Bartlett Deep: American Journal Science, v. 239, p. 128-
147.
, and PARKER, F. L., 1947, Bulimina and related foraminif-
eral genera: U S . Geological Survey Professional Paper 210-D,
p. 55-176, pls. 15-30.
DAULTRY,S., 1976, Principal Components Analysis: Geo Ab-
stracts Ltd., University of East Anglia, Norwich, 50 p.
DAVIS,J. C., 1973, Statistics and Data Analysis in Geology: John
Wiley and Sons, 550 p.
DONN,W. L., 1975, Meteorology, McGraw-Hill Book Company,
518 p.
ELLIS,B. F. and MESSINA, A. R., 1940, Catalogue of Foraminifera.
Special Publication. The American Museum of Natural History,
(supplements post-1940).
EMILIANI, C., 1971, Paleotemperature variations across the Plio-
Pleistocene boundary: Science, v. 1977, p. 60-62.
PLIOCENE-PLEISTOCENE IN ITALY
EMILIANI,J. A., MAYEDA, T., and SELLI,R., 1961, Paleotemper-
ature analysis of the Plio-Pleistocene section at Le Castella,
Calabria, southern Italy: Geological Society America Bulletin,
p. 679-688.
FLINT,J. M., 1899, Recent Foraminifera (A descriptive catalogue
of specimens dredged by the U.S. Fish Commission steamer
Albatross): Annual Report (1897), United States National Mu-
seum, p. 249-349.
FORNASINI, C., 1900, Intorno ad alcuni esemplaridi foraminiferi
Adriatici: R. Accad. Sci. 1st. Bologna, Mem., Bologna, Italia,
1900, ser. 5 , tom0 8, p. 393, in Ellis, B., and Messina, A., 1940,
Catalogue of Foraminifera: American Museum Natural History
(supplements, post- 1940).
, 1901, Boll. SOC.Geol. Ital., v. 20, p. 174, plate 1, in Ellis,
B., and Messina, A., 1940, Catalogue of Foraminifera: Amer-
ican Museum Natural History (supplements, post-1940).
GIGNOUX, M., 1910, Sur la classification du Pliocene et du Qua-
ternaire dans 1’Italie du Sud. Comptes-Rendus de l’ Acadtmie
des Sciences S i n e D, Volume 150, pp. 841-844.
GRADSTEIN, F. M., 1970, Foraminifera from the type Sicilian at
Ficarazzi, Sicily (Lower Pleistocene): Koninklijke Nederlandse
Akademie van Wetenschappen, Series B, v. 73, no. 4, p. 1-29.
HAQ, B. U., BERGGREN, W. A., and VAN COUVERING, J. A.,
1977, Corrected age of the Pliocene/Pleistocene boundary: Na-
ture, v. 269, no. 6, p. 483-488.
HAQ, B. U., and LOHMANN, G. P., 1976, Early Cenozoic calcar-
eous nannoplankton biogeography of the Atlantic Ocean: Ma-
rine Micropaleontology, v. l , no. 2, p. 119-194.
HERON-ALLEN, E., and EARLAND, A., 1913, The foraminifera of
the Clare Island District, Co. Mayo, Ireland: Royal Irish Acad-
emy Proceedings, v. 31, pt. 64,p. 1-188.
HOFKER,J., 1932, Notizen ueber die Foraminiferen des Golfes
von Neapel. 111. Die Foraminiferenfauna der Ammontatura:
Stazione Zoologica di Napoli Publicazioni, v. 12, fasc. 1, p. 61-
144.
, 1951, The foraminifera of the Siboga Expedition, Part 3:
Monograph 4a, I-XII, p. 1-513.
HOOPER,K., 1969, A re-evaluation of eastern Mediterranean fo-
raminifera using factor-vector analysis: Contributions from the
Cushman Foundation for Foraminiferal Research, v. 20, no. 4,
p. 147-151.
IACCARINO, S., 1975, Planktonic and significant benthonic fora-
minifera of the proposed Plio-Pleistocene boundary type-sec-
tion of Le Castella: L’Atemeo Parmense, Acta Naturalia, v.
IO, p. 449-465.
IMBRIE,J., and KIPP, N. G., 1971, A new micropaleontological
method for quantitative paleoclimatology: Application to a Late
Pleistocene Caribbean core, in Turekian, K. K., ed., The Late
Cenozoic Glacial Ages, Yale University Press, p. 71-181.
KLOVAN,J. E., and IMBRIE,J., 1971, An algorithm and FOR-
TRAN-IV program for large-scale Q-mode factor analysis and
calculation of factor scores: Mathematical Geology, v. 3, p. 61-
77.
LACOMBE, H., and TCHERNIA, P., 1972, Caracteres Hydrologiques
et Circulation des Eaux en Mediterrante, in Stanley, D. J., ed.,
The Mediterranean Sea: A Natural Sedimentation Laboratory:
Stroudsburg, Pennsylvania, Dowden, Hutchinson and Ross,
Inc., p. 149-176.
LALICKER, C. G., 1935, Two new foraminifera of the genus Tex-
tularia: Smithsonian Institution Miscellaneous Collections,
Washington, D.C., v. 91, no. 22, pl. 1.
LAMB,J. L., and BEARD,J. H., 1972, Late Neogene planktonic
foraminifera in the Caribbean Gulf of Mexico and Italian strato-
15
BREMER AND OTHERS
types: University of Kansas Paleontology Contribution Article
57 (Protozoa 8), p. 1-67.
LINNAEUS, C., 1791, Sysrema Naturae. Ed B, aucta reformata.
Cura Jo. Frid. Gmelin. Lipsiae, impensis G. E. Beer, Tom. 1,
pt. 6 (Vermes), pp. 3,021-3,910.
LOEBLICH, A. R., JR., and TAPPAN,H., 1964, Sarcodina, chiefly
“Thecamoebians” and Foraminiferida, in Moore, R. C., ed.,
Treatise on Invertebrate Paleontology, Protista Z, pt. c, Kansas
Univ. Press.
LOHMANN, G. P., 1978, Abyssal benthonic foraminifera as hydro-
graphic indicators in the western South Atlantic Ocean: Journal
of Foraminiferal Research, v. 8, no. 1, p. 6-34.
LONGINELLI, A., 1956, Foraminiferi del Calbriano Piacenziano:
Palaeontolographia Italica, v. 49, p. 1-1 16.
MARCHETTI, D. B., 1975, Preliminary palynological data on the
proposed Plio-Pleistocene boundary type-section of Le Castel-
la: L’Ateneo Parmense, Acta Naturalia, v. 1 1 , p. 457-485.
MARKS,P., 1951, A revision of the smaller foraminifera from the
Miocene of the Vienna Basin: Cushman Foundation Fora-
miniferal Research Contribution, v. 2, pt. 2, p. 33-73.
MURRAY, J. W., 1971, An Atlas of British Recent Foraminiferids:
American Elsevier Publishing Company, New York, 244 p.
NORVANG, A., 1958, Zslandiella n.g. and Cassidulina d’orbigny.
Videnskabelige Meddelelser, Dansk Naturhistorisk Forening,
Bind 120, pp. 25-41.
ORBIGNY, D., A. D., 1826, Tableau mtthodique de la classe des
Ctphalopodes: Annales des Sciences Naturelles, v. 7, p. 245-
3 14.
-, 1839, Foraminiferes, in De la Sagra, Historia Fiscia, Pol-
itica y Natural de la Isla de Cuba, 2 vol. 1837-1842, p. 1-224.
-, 1846, Foraminieres fossiles du Bassin Tertiaire de Vienne:
Pans, 312 p.
, 1852, Du Prodrome de paltontologie stratigraphique uni-
verselle des animaux mollusques et rayonnes. Paris, Masson,
Vol. 3, pp. 1-196.
PARKER, F. L., 1954, Distribution of the Foraminifera in the north-
eastern Gulf of Mexico: Museum Comparative Zoology Bulle-
tin, v. 1 1 1 , no. 10, p. 453-588.
, 1958, Eastern Mediterranean foraminifera: Reports of the
Swedish Deep-sea Expedition, v. 8, p. 219-283.
PARKER, W. K., and JONES,T. R., 1865, On some Foraminifera
from the North Atlantic and Arctic Oceans, including Davis
Straits and Baflin’s Bay: Philosophical Transactions, p. 325-
441.
PFLUM,C. E., and FRERICHS, W. E., 1976, Gulf of Mexico Deep-
Water Foraminifera: Cushman Laboratory Foraminiferal Re-
search, Special Publication no. 14, p. 7-125.
PHLEGER, F. B., and PARKER, F. L., 1951, Ecology offoraminifera,
northwest Gulf of Mexico: Geological Society of America
Memoir 46, pt. 2, p. 1-64.
PHLEGER, F. B., PARKER, F. L., and PEIRSON,J. F., 1953, North
Atlantic foraminifera: Reports of the Swedish Deep-sea Ex-
pedition, v. 7, p. 1-122.
POLLAK, M. J., 1951, The sources of the deep water of the Eastern
Mediterranean Sea: Journal Marine Research, v. 10, no. 1, p.
129-152.
PUJOS-LAMY, A., 1973, Rtpartition des Biocoenoses de Foramin-
16
&res Benthiques sur le Plateau Continental du Golfe de Gas-
cogne a I’ouest de l’embouchure de la Gironde. Revista Espafi-
ola de Micropaleontologia, vol. IV, num. 2, pp. 141-156.
REUSS,A. E., 1850, Neue Foraminiferen aus den Schichten des
osterreichischen Tertiarbeckens: Denkschriften der Akademie
der Wissenshaften (Wien); Mathematisch-Naturwissen-
schaftlich Classe, v. 1 , p. 365-390.
RIO, D., 1974, Remarks on Late Pliocene-Early Pleistocene cal-
careous nannofossil stratigraphy in Italy: L’Ateneo Parmense,
Acta Naturalia, v. 10, p. 409-449.
RYAN,W. B., 1972, Stratigraphy of Late Quaternary sediments in
the eastern Mediterranean, in Stanley, D. J., ed., The Medi-
terranean Sea: A Natural Sedimentation Laboratory: Strouds-
burg, Pennsylvania, Dowden, Hutchinson and Ross, Inc., p.
147- 176.
SCHROETER, J. G., 1783, Einleitung in die conchylienkenntniss
nach Linne, v. 1 , p. 20, pl. 1 , fig. 2, in Ellis, B., and Messina,
A., 1940, Catalogue of Foraminifera: American Museum Nat-
ural History (supplements, post- 1940).
SEGUENZA, G., 1862, Prime ricerche intorno ou rhizopodi fossili
delle Argille Pleistoceniche dei dentorni de Catania: Atti
dell’Accademia Gioenia di Scienze Naturali in Catania, ser. 2,
V. 18, p. 85-126.
SILVESTRI, A,, 1893, Su di aleuni foraminifen del Mare Jonio: Ac-
cad. Pont. Muovi Lincei, Mem., Roma, Italia, v. 9, p. 200, in
Ellis, B., and Messina, A., 1940, Catalogue of Foraminifera:
American Museum Natural History (supplements, post-1940).
, 1896, Foraminiferi Pliocenici della Provincia di Siena. Part
1: Pontificia Academia Scientiarum, Memorie, v. 12, p. 1-204.
-, 1898, Foraminiferi pliocenici della Provincia di Siena: Parte
11: Accad. Pont. Nuovi Lincei, Mem., Roma, Italia, v. 15, p.
329, in Ellis, B., and Messina, A., 1940, Catalogue of Fora-
minifera: American Museum Natural History (supplements,
post- 1940).
-, 1904, Ricerche strutturali su alcune forme dei Trubi di
Bonfornello (Palermo): Pontificia Accademia Scientiarium,
Memorie, v. 22, p. 235-276.
SVERDRUP, H. U., JOHNSON, M. W., and FLEMING, F. H., 1942,
The Oceans, their physics, chemistry and general biology.
Prentice-Hall, N.J., 1,087 p.
TODD,R., 1958, Foraminifera from western Mediterranean deep-
sea cores: Reports of the Swedish Deep-sea Expedition, v. 8,
no. 3, p. 169-216.
TREWARTHA, G. T., 1968, An Introduction to Climate. McGraw-
Hill Book Company, 408 p.
VANLOON,H., 1972, Wind in the Southern Hemisphere: Meteo-
rological Monographs, v. 13, no. 35, p. 87-99.
WALKER,G., and JACOB,E., 1798, Adams’ Essays on the micro-
scope. Ed. Z, Dillon and Keating, London, England, p. 642, in
Ellis, B., and Messina, A., 1940, Catalogue of Foraminifera:
American Museum Natural History (supplements, post- 1940).
WILLIAMSON, W. C., 1858, On the Recent foraminifera of Great
Britain: Royal Society of London, England, p. 77, in Ellis, B.,
and Messina, A,, 1940, Catalogue of Foraminifera: American
Museum Natural History (supplements, post-1940).
Revised Ms accepted August 1979
 PLIOCENE-PLEISTOCENE IN ITALY

APPENDIX 1 Bulimina marginata d’orbigny, 1826

PI. 1, Fig. 7
TAXONOMIC
NOTES
Bulimina marginata D’ORBIGNY, 1826, p. 269, no. 4, pl. 12, figs.
In this section we present synonymies and a few brief discussions 10-12. BRADY,1884, pl. 51, figs. 3-5. CUSHMAN AND PARKER,
of those taxa which constitute 5% or more of the total fauna in at 1947, p. 119, pl. 28, figs. 5 , 6 (extensive synonymy). PHLEGER
least one sample. Because the paleoenvironmental reconstruction A N D PARKER,1951, p. 16, pl. 7, figs. 27, 28. PHLEGER AND
is based on comparison of the Le Castella faunas and the modern OTHERS,1953, p. 262, pl. 6, figs. 25, 26. PARKER,1958, p. 262,
faunas of the eastern Mediterranean described by Parker (1958), the pl. 2, fig. 23. BELFORD,1966, p. 555, pl. 5 , figs. 4, 5. MURRAY,
taxonomic concepts are, wherever possible, the same. 1971, p. 119, pl. 49, figs. 1-7. COLOM,1974, p. 117, text figs.
16: d’-h’, k’-0’.
Family TEXTULARIIDAE
Bulimina costata d’Orbigny, 1826
Textularia calva, Lalicker, 1935 PI. 1, Fig. 8
PI. 1, Fig. 1
Bulimina costata, D’ORBIGNY,1826, p. 269, no. 1. D’ORBIGNY,
Textularia calva, LALICKER,1935, p. 1, pl. 1, figs. 1-2. PARKER, 1852, p. 194. FORNASINI, 1901, p. 174, text fig. 1. CUSHMAN
1958, p. 254, pl. 1, figs. 3, 4. A N D PARKER, 1947, p. 115, pl. 27, figs. 2, 3. PARKER,1958, p.
261, pl. 2, figs. 19, 20. MURRAY, 1971, p. 115, pl. 47, figs. 1-7.
Family MILIOLIDAE
Bulimina exilis Brady, 1884
Sigmoilopsis schlumbergeri (Silvestri), 1884 PI. 1, Fig. 10
PI. 1, Figs. 2, 3
Bulimina elegans d’Orbigny var. exilis BRADY,1884, p. 399, pl. 50,
Planispirina celata BRADY,1884, p. 192, pl. 8, figs. 1-4, figs. 5 , 6.
(non Spiroloculina celata Costa). Bulimina exilis Brady. PHLEGER A N D OTHERS, 1953, p. 32, pl. 6, fig.
Sigmoilina schlumbergeri SILVESTRI,1904, p. 267. PARKER,1958, 24. PARKER,1958, p. 261, pl. 2, fig. 29.
p. 256.
Sigmoilopsis schlumbergeri (Silvestri). BARKER,1960, p. 16, pl. 8,
Bulimina gibba Fornasini, 1901
figs. 1-4.
PI. I , Fig. 9
Family NODOSARIIADAE Bulimina gibba FORNASINI, 1901, p. 378, pl. 0, figs. 32-34. PARKER,
1958, p. 261, pl. 2, fig. 29.
Amphicoryna scalaris (Batsch, 1791)
PI. 1, Fig. 4
Globobulimina affinis d’Orbigny, 1939
Nautilus scalaris BATSCH,1791, pl. 2, figs. 4a, b. PI. 1, Fig. 5
Lagenodosaria scalaris (Batsch). TODD,1958, p. 258, pl. 1, figs. 32,
Bulimina afinis D’ORBIGNY, 1939, p. 105, pl. 2, figs. 25,26. PARKER,
33.
1958, P. 262, PL. 2, FIGS.24, 25.
Amphicoryna scalaris (Batsch). BARKER,1960, pl. 63, figs. 28-31.
BERGGREN AND OTHERS,1976, p. 209, pl. 2, figs. 3, 4.
FAMILY
BOLIVINITIDAE
Family BULIMINIDAE Brizalina attica (Parker), 1958
PI. 1, Fig. 13
Bulimina aculeata d’orbigny, 1826
PI. 1, Fig. 6 Bolivina attica PARKER,1958, p. 259, pl. 2, figs. 12-14.
Bulimina aculeata D’ORBIGNY, 1826, p. 269, no. 7. BRADY,1884,
pl. 51, figs. 7-9. CUSHMAN AND PARKER, 1947, p. 120, pl. 28, Brizalina catanensis (Seguenza), 1862
figs. 8-11. PHLEGER AND OTHERS, 1953, p. 32, PI. 6, figs. 21, PI. 1, Fig. 15
22. PARKER,1958, p. 261, pl. 2, figs. 17, 18. BELFORD,1966, p. Bolivina catanensis SEGUENZA, 1862, p. 29, pl. 2, figs. 3a, b. PAR-
58, pl. 5 , figs. 1-3. COLOM, 1974, p. 115, text fig. 16:g. BERG- KER,1958, pl. 2, figs. 10, 11.
GREN AND HAQ, 1976, p. 108, pl. 4, figs. 1-5.
Remarks. Distinction between this species and the apparently re-
lated form B . marginata d’Orbigny has been made on the basis of Brizalina difformis (Williamson), 1858
the undercut chamber margins, particularly in the early stages, in PI. 1, Fig. 12
B . marginata and the greater development of apically directed Textularia variabilis Williamson var. difformis, WILLIAMSON, 1858,
spines in B . aculeata (F. Parker, personal communication; Belford, p. 77, pl. 6, figs. 166, 167.
1966). Many individuals at Le Castella possess the attributes of Bolivina difformis (Williamson). PARKER, 1958, p. 260, pl. 2, fig. 9.
both. It is possible that they are intergrades between two species,
that they represent intergrades between two end-members of one Brizalina dilatata (Reus), 1850
species or they are the megalospheric and microspheric forms of PI. 1, Fig. 14
one species. The problem needs further study. For the purposes of
this study, the two taxa were lumped. We illustrate the end-mem- Bolivina dilatata REUS, 1850, p. 381, pl. 48, figs. 5a-c. CUSHMAN,
bers in the plates. 1937, p. 78, PI. 9, figs. 17-20.

17
BREMER AND OTHERS
Bolivina dilatata Reuss var. dilitntissima SILVESTRI,
1893, p. 100.
PARKER,1958, p. 260, pl. 2, figs. 5-7.
Brizalina alata (Seguenza), 1862
PI. 1, Fig. 11
Vulvulina d a t a SEGUENZA, 1862, p. 115, pl. 2, figs. 5, Sa.
Bolivina beyrichi var. alata Seguenza. BRADY,1884, p. 442, PI. 53,
figs. 2-4.
Bolivina alata (Seguenza). CUSHMAN,1937, p. 106, pl. 13, figs. 3-
11. PHLEGERA N D PARKER,1951, p. 12, pl. 6, fig. 11. COLOM,
1974, p. 119, text fig. 17: 11-12.
Loxostomum alatum (Seguenza). PARKER,1958, p. 262, pl. 2, figs.
30-32.
Brizalina alata (Seguenza). BELFORD,1966, p. 24, pl. 1, fig. 12.
BERGGREN AND HAQ, 1976, p. 75-78, pl. 3, fig. 1.
Family SPHAEROIDINIDAE
Sphaeroidina bulloides d’Orbigny , 1826
PI. 2, Figs. 5, 6
Sphaeroidina bulloides D’ORBIGNY,1826, p. 267, no. 1, Modtles,
no. 65. PARKER A N D JONES,1865, pl. 2, fig. 58 (figure after
d’orbigny’s Modkle). GRADSTEIN,1970, p. 23, figs. 3a-d.
PLATE 1
1 Textularia calva Lalicker. Sample 36. x 113.
2, 3 Sigmoilopsis schlumbergeri (Silvestri). 2. Apertural view,
sample 40, ~ 6 2 3.
. Front view, sample 59, ~ 7 2 .
4 Amphicoryna scalaris (Batsch). Sample 47, X 157.
5 Globobulimina af3nis (d’orbigny). Sample 28, X 136.
6 Bulimina aculeata d’Orbigny. Sample 55, X 110.
7 Bulimina marginata d’Orbigny. Sample 60, x 144.
8 Bulimina costafa d’Orbigny. Sample 60, X210.
PLATE 2
1 Uvigerina auberiana d’Orbigny. Sample 45, x 153.
2, 3 Uvigerina peregrina Cushman. 2. Sample 54, X I O O . 3.
Sample 3 1, x 109.
4 Trifarina angulosa (Williamson). Sample 52, X 127.
5 , 6 Sphaeroidina bulloides d’orbigny. 5. Sample 36, x 124. 6.
Sample 59, x 118.
7, 8 Globocassidulina subglobosa Brady. 7. Sample 32, X 122.
8. Sample 45, x 112.
PLATE 3
1, 2 Cibicides boueanus (d’orbigny). 1. Umbilical view, sample
51, x190. 2. Spiral view, sample 41, ~ 1 8 6 .
3, 4 Pullenia bulloides (d’orbigny). 3. Apertural view, sample
33, x80. 4. Side view, sample 46, ~ 1 0 4 .
5-7 Planulina arimenensis d’Orbigny. 5 . Umbilical view, sam-
ple 59, x95. 6. Apertural view, sample 58, X82. 7. Spiral
view, sample 44, ~ 6 0 .
18
Family UVIGERINIDAE
Uvigerina auberiana d’Orbigny , 1829
PI. 2, Fig. 1
Uvigerina auberiana D’ORBIGNY, 1829, p. 106, pl. 2, figs. 23, 24.
BERGGREN A N D OTHERS, 1976, p. 211, pl. 3, figs. 1-7.
Uvigerina peregrina Cushman, 1923
PI. 2, Figs. 2, 3
Uvigerina pygmaea d’orbigny. BRADY,1884, pl. 74; figs. 11, 12
(non d’orbigny). FLINT,1899, p. 320, pl. 68, fig. 2.
Uvigerina peregrina CUSHMAN,1923, p. 166, pl. 42, figs. 7-10.
PHLEGER A N D PARKER,1951, p. 18, pl. 8, figs. 22, 24-26. PAR-
KER, 1958, p. 262, pl. 2, figs. 37, 38. MURRAY, 1971, p. 121, pl.
50, figs. 1-7. PFLUM A N D OTHERS, 1976, p. 25, pl. 8, figs. 2, 3.
Uvigerina mediterranea HOFKER,1932, p. 118, fig. 32a-f. LONGI-
NELLI,1956, p. 163, pl. 23, figs. 20, 25.
Trifarina angulosa (Williamson), 1858
PI. 2, Fig. 4
Uvigerina angulosa WILLIAMSON, 1858, p. 67, PI. 5, fig. 140. BRA-
DY, 1884, pl. 74, figs. 15, 16.
Angulogerina angulosa (Williamson). PARKER,1958, p. 259, pl. 21,
figs. 1, 2.
9 Bulimina gibba Fornasini. Sample 8, X 136.
10 Bulimina exilis Brady. Sample 18, x 136.
11 Brizalina alata (Seguenza). Sample 9, ~ 3 9 .
12 Brizulina difformis (Williamson). Sample 26, X 119.
13 Brizalina nttica (Parker). Sample 29, x 106.
14 Brizalina dilitata (Reuss). Sample 50, x 120.
15 Brizalina catanensis (Seguenza). Sample 51, ~ 8 6 .
9, 10 Cassidulina laevigata d’orbigny. 9. Sample 55, X56. 10.
Sample 51, ~ 9 5 .
11, 12 Cassidulina crassa d’Orbigny. 11. Sample 19, ~ 2 0 9 .12.
Sample 19, x157.
13-15 Epistomineh evax Bandy. 13. Umbilical view, sample 36,
x 144. 14. Apertural view, sample 26, x69. 15. Spiral view,
sample 26, x 106.
8 Hyalinea balthica (Gmelin). Sample 35, X 120.
9-11 Cibicides lobatulus (Walker and Jacob). 9. Umbilical view,
sample 38, x49. 10. Apertural view, sample 38, ~ 8 2 11..
Spiral view, sample 38, ~ 8 9 .
12-14 Cibicidoides floridanus (Cushman). 12. Umbilical view,
sample 59, ~ 9 0 13.
. Apertural view, sample 44, X46. 14.
Spiral view, sample 39, ~ 8 0 .
PLIOCENE-PLEISTOCENE IN ITALY

19
BREMER AND OTHERS

20
PLIOCENE-PLEISTOCENE IN ITALY

21
BREMER A N D OTHERS

22
 PLIOCENE-PLEISTOCENE IN ITALY

Trifarina angulosa (Williamson). LOEBLICH A N D TAPPAN,1964, p. Hyalinea balthica (Gmelin), 1791

571, pl. 450, figs. 1-3 (paratypes figures; lectotype 2F3576 and
paratypes 2F3575 designated). MURRAY, 1971, p. 123, pl. 51,
figs. 1-6. COLOM,1974, p. 123, text figs. 19, q-t.
Family DISCORBIDAE
Epistominella evax, Bandy, 1953
PI. 2, Figs. 13-15
Epistominella evax BANDY,1953, pl. 23, p. 179.
Valvulineria bradyana (Fornasini), 1846
?Discorbina bradyana FORNASINI, 1900, p. 393, text fig. 43.
Valvulineria complanata (d’orbigny). MARKS,1951, p. 64, pl. 6,
figs. 13a-c. TODD,1958, p. 196. PARKER,1958, p. 268, 269, pl.
3, figs. 42-44.
Remarks. Le Castella specimens agree well with the type illus-
tration of bradyana, with specimens referred to this species by
Mediterranean authors, but also with specimens identified as V.
complanata in the collections of F. L. Parker. However, the type
illustrations of complanata differ significantly from that of bradyana
and it is not clear whether the two forms are conspecific. Pending
classification of this, the name bradyana is used here in preference
to complanata.
Gavelinopsis praegeri (Heron-Allen and Earland),
1913
Discorbina praegeri HERON-ALLEN AND EARLAND, 1913, p. 122,
PI. 10, figs. 8-10.
Gavelinopsis praegeri (Heron-Allen and Earland). TODD, 1958, p.
197. PARKER, 1958, p. 264, pl. 3, figs. 24,25. GRADSTEIN, 1970,
p. 15. MURRAY, 1971, p. 133, pl. 55, figs. 1-5.
Family CIBICIDIDAE
Cibicides boueanus (d’orbigny), 1846
PI. 3, Figs. 1, 2
Truncatulina boueana D’ORBIGNY,
1846, p. 169, pl. 9, figs. 24-26.
Cibicides lobatulus (Walker and Jacob), 1798
PI. 3, Figs. 9-11
Nautilus lobatulus WALKERAND JACOB,1798, p. 642, pl. 14, fig.
36.
Cibicides lobatulus (Walker and Jacob). CUSHMAN,1931, p. 118,
pl. 21, fig. 3. PARKER,1958, p. 275. GRADSTEIN,
1970, p. 13.
MURRAY,1971, p. 175, pl. 73, figs. 1-7.
PI. 3, Fig. 8
“Ammonshorner der ostsee,” Schroeter, 1782. SCHROETER, 1783,
p. 20, PI. 1, fig. 2.
Nautilus balthicus GMELIN,1791, in Linnaeus, Systema Naturae,
13th ed. Leipzig, p. 3370.
Operculina annomoides BRADY,1884 (non Gromovius), p. 745, pl.
112, figs. 1, 2.
Anomalina balthica (Schroeter). CUSHMAN, 1931, p. 108, pl. 19, fig.
3.
Hyalinea balthica (Schroeter). HOFKER,1951, p. 508. PARKER,
1958, p. 275, pl. 4, fig. 39. LOEBLICH A N D TAPPAN,1964, p.
686, fig. 552, no. 2, 3. BELFORD,1966, p. 124, pl. 14, figs. 1-8.
GRADSTEIN, 1970, p. 17, pl. 1, figs. 15, 16. MURRAY, 1971, p.
173, pl. 72, figs. 5-8. COLOM,1974, p. 151, fig. 37.
Planulina arimenensis d’orbigny, 1826
PI. 3, Figs. 5-7
Planulina arimenensis D’ORBIGNY, 1826, p. 280, no. 1, pl. 14, figs.
1-3. BRADY,1884, pl. 93, figs. 10, 11. PHLEGER AND OTHERS,
1953, p. 48, pl. 11, figs. 3, 4. PARKER,1958, p. 276, pl. 4, fig.
43. LOEBLICH AND TAPPAN,1964, p. 686, fig. 552, la-c. BERG-
GREN AND HAQ, 1976, 107, pl. 2, fig. 11.
Family CASSIDULINIDAE
Cassidulina crassa d’Orbigny, 1839
PI. 2, Figs. 11, 12
Cassidulina crassa D’ORBIGNY, 1839, p. 56, pl. 7, figs. 18-20. PHLE-
GER AND PARKER,1951, p. 12, pl. 14, fig. 4a, b. PARKER, 1958,
p. 44, pl. 10, fig. 1. BARKER,
1960, pl. 54, figs. 4,5. GRADSTEIN,
1970, p. 12.
Globocassidulina crassa (d’orbigny) BELFORD,1966, pl. 26, figs.
5-9.
Cassidulina laevigata d’Orbigny, 1826
PI. 2, Figs. 9, 10
Cassidulina laevigata D’ORBIGNY,1826, p. 282, pl. 15, figs., 4, 5.
BELFORD,1966, p. 138, pl. 24, figs. 1-4; text figs. 16: 1-2.
Cassidulina laevigata d’Orbigny var. carinata SILVESTRI, 1896, p.
104, pl. 2, fig. 10 (C. laevigata in expl. of plate).
Cassidulina carinata Silvestri. PHLEGER AND OTHERS,1953, p. 44,
PI. 9, figs. 32-37. PARKER,1958, p. 271, pl. 4, figs. 15. MURRAY,
1971, p. 187, PI. 78, figs. 1-5.
Remarks. The taxonomy of carinate vs. non-carinate cassidulinids
is complex. We follow Belford (1966) who, based on morphologic

PLATE 4
1-3 Gyroidina umbonata (Silvestri). 1. Umbilical view, sample 7-9Oridorsalis tener (Brady). 7. Umbilical view, sample 14,
39, x 120. 2. Apertural view, sample 41, x 148. 3. Spiral x 113. 8. Apertural view, sample 29, ~ 9 6 9.
. Spiral view,
view, sample 41, x 199. sample 14, x94.
4-6 Gyroidinoides neosoldanii (Brotzen). 4. Umbilical view, 10, 11 Melonis barleeanus (Williamson). 10. Sample 59, X 105. 11.
sample 36, x110. 5. Apertural view, sample 37, x110. 6. Apertural view, sample 49, x 107.
Sprial view, sample 45, x 126. 12, 13 Melonis soldanii (d’orbigny). 12. Apertural view, sample
39, x95. 13. Sample 40, x 152.

23
BREMER AND OTHERS

studies by Nprvang (1958) considered the carinate (carinata) and Family OSANGULARIIDAE
non-carinate or weakly carinate (laevigata) forms as taxonomically
identical. Le Castella individuals agree well with those identified as Gyroidinoides neosoldanii (Brotzen), 1936
C. carinata Silvestri in the collections of F. L. Parker. They are PI. 4, Figs. 4-6
peripherally compressed with slight to moderate umbonal inflation, Gyroidina neosoldanii, BROTZEN,1936, p. 158.
generally subcircular in outline (although some individuals exhibit Gyroidina cf. G . neosoldanii Brotzen. PARKER,1958, p. 265, pl. 3,
an undulating, petaloid periphery). figs. 13-18.
Gyroidinoides neosoldanii Brotzen. BERGGREN A N D OTHERS, 1976,
Globocassidulina subglobosa Brady, 1881 p. 217, pl. 5 , figs. 6-8.
PI. 2, Figs. 7, 8
Cassidulina subglobosa Brady. BRADY,1884, p. 430, pl. 54, fig. 17.
Family ANOMALINIDAE
PHLEGER A N D OTHERS,1953, p. 45, pl. 10, fig. 4. PARKER,
1958,
p. 272, pl. 4, fig. 13. Melonis barleeanus (Williamson), 1858
Globocassidulina subglobosa (Brady). BELFORD,1966, p. 149, pl. PI. 4, Figs. 10, 11
25, figs. 11-16. LOHMANN, 1978, pl. 2, figs. 8, 9.
Nonionina barleeana WILLIAMSON, 1858, p. 32, pf. 3, figs. 68, 69.
Nonion barleeanum (Williamson). PARKER, 1958, p. 258, pl. 1, figs.
Family NONIONIDAE
36, 37.
Pullenia bulloides (d’orbigny), 1826 Melonis barleeanus (Williamson). BERGGREN A N D HAQ, 1976, p.
PI. 3, Figs. 3, 4 105, pl. 1, figs. 19, 20. BERGGRENAND OTHERS, 1976, p. 220,
pl. 4, fig. 13, 14.
Nonionina bulloides D’ORBIGNY, 1826, p. 293. D’ORBIGNY,1846, Remarks. Melonis barleeanus is more compressed and less deep-
p. 107, pl. 5, figs. 9, 10. ly umbilicate than Melonis soldanii. Because the two taxa inter-
Pullenia sphaeroides (d’orbigny). BRADY,1884, p. 615, pl. 84, figs. grade at Le Castella, they were lumped. We illustrate the two end-
12, 13. members in the plates.
Pullenia bulloides (d’orbigny). PARKER, 1954, p. 538, pl. 11, fig. 17.
BERGGREN A N D OTHERS, 1976, v. 1, p. 217, pl. 5 , figs. 6-8.
Melonis soldanii (d’orbigny), 1846
Florilus boueanum (d’orbigny), 1846 PI. 4, Figs. 12, 13
Nonionina boueana, D’ORBIGNY, 1846, p. 108, PI. 5, figs. 11, 12. Nonionina soldanii D’ORBIGNY, 1846, p. 109, figs. 15, 16.
Nonion boueana (d’orbigny). CUSHMAN, 1939, p. 12, pl. 3, figs. 7, Nonion soldanii (d’orbigny). CUSHMAN, 1939, p. 13, figs. 10, 11,
8. pl. 4, fig. 23.
Florilus boueanum (d’orbigny). BERGGREN A N D HAQ, 1976, p. 107,
pl. 3, figs. 15-18.
Cibicidoides floridanus (Cushman), 1918
Family ALABAMINIDAE PI. 3, Figs. 12-14

Gyroidina umbonata (Silvestri), 1898 Truncatulinafloridana CUSHMAN, 1918, p. 62, pl. 19, fig. 2.
PI. 4, Figs. 1-3 Cibicides aff. floridanus (Cushman). PHLEGER A N D PARKER, 1951,
p. 30, pl. 16, figs. 1-4. PARKER,1958, p. 274, pl. 4, figs. 36-38.
Rotalia soldanii d’Orbigny var. umbonata, SILVESTRI, 1898, p. 329, Cibicidoides Boridana (Cushman). BERGGREN A N D HAQ, 1976, p.
pl. 6, figs. 14a-c. 107, pl. 3, figs. 13, 14.
Gyroidina umbonata (Silvestri). PARKER, 1958, p. 197, pl. 1, fig. 18. Remarks. Phleger and Parker (1951) called attention to the highly
variable nature of this species. Parker (1958) noted the apparent
Oridorsalis tener (Brady), 1884 relationships of this form to the ungerianus-pseudocrngerianus
PI. 4, Figs. 7-9 group. In fact, an examination of a large number of specimens in F.
Oridorsalis tener (Brady). BERGGREN A N D OTHERS,
1976, pl. 4, figs. L. Parker’s collections reveal that she also included under the name
6-8. LOHMANN, 1978, pl. 4, figs. 5-7. floridanus specimens referable to C. mediocris Finlay (= C . pseudo-
Oridorsalis tener tener (Brady). PFLUMA N D OTHERS,1976, p. 218, ungerianus Cushman, in part, not the holotype; see Belford, 1966;
pl. 6, figs. 2-4. Berggren and others, 1976).
Remarks. We follow the species concept of Lohmann (1978) who The characteristics which distinguish C.floridanus and mediocris
points out that this species of Oridorsalis is characterized by the were discussed by Parker (1958, p. 275); she combined these two
curved nature of the sutures on the spiral side, the equal size of the taxa because they intergrade. Similar intergradation is seen through-
chambers in the final whorl, and the compressed cross-sectional out the section at Le Castella, and for this reason, the two taxa
outline. were combined.

24
 PLIOCENE-PLEISTOCENE I N ITALY

APPENDIX 2
Percentages of species of benthonic foraminifera from samples (sample numbers across the top of table) of the Le Castella.

22 21 20 19 18 17 16 15 14 13 12 II IO 9 8 7 6 5

Ammonia beccarii 1 1 1 1 3
Asterigerinita mammila
Bigeneria nodosaria 1 1
Brizalina albatrossi 1 1 2 1 1 1
Brizalina attica 2 34 51 3 2 63 45 8 13 53 11
Brizalina catanensis 9 7 1 1 2 1 14 I 23 1 18 1
Brizalina difformis 3 14
Brizalina dilitata 3
Brizalina pseudoplicata 3
Brizalina alata
Bulimina aculeata + marginata 5 2 6 8 1 2 9 1 19
Bulimina costata 4 4 1 6 1 4 3 1 8 7 2 1 1 4 1 3 2 0 8 1
Bulimina exilis 1 44 36 1 9 14 1 4 86 8 2
Bulimina gibba 1 3 1 1 I 1 3
Cibicides boueanus 2 1 1 5 4 2 2 1
Cassidulina crassa
Cassidulina laevigata 4 7 8 6 2 1 1 0 1 3 9 1 1 1 5 1 22
Globocassidulina subglobosa 6 1 4 4 1 1 1 1 I 1 4 1 6
Cibicides lotatulus 1 1 2 1 9
Cibicidoides floridanus 33 36 16 14 1 1 39 29 7 43 1 53 11 43 9 1
Eggerella bradyi 1 1
Elphidium crispum
Epistominella evax
Epistominella rugosa convexa 1
Florilus boueanum I 1 7
Gavelinopsis praegeri 1 1
Globobulimina affinis 2 1 2 1 1 2 1 2 1
Globobulimina pseudospinescens 1 4 3 1 2
Gyroidina altiformis
Gyroidina umbonata 3 3 1 1 1 1 1 1
Gyroidinoides neosoldanii 1 1 3 2 3 2 1 1 1
Hanzawaia rhodiensis
Hoglundina elegans 1 6
Hyalinea balthica
Amphicoryna scalaris
Melonis barleanus 1 1 3 3 3 1 5 9 1 13 11 3 5
Miliolinella circularis 2
Neoconorbina terquemi 1 1 1 1 2 1
Oridorsalis tener 2 1 2 2 1 2 7 2 1 3
Planorbulina mediterranensis
Planulina arimenensis 1 8 1 2 5 5 1 4 5 1 3
Pseudoclavulina crustata 1 1
Pullenia bulloides 1 2 2 1 1 3 2 1 1 1 I
Pullenia quinqueloba 1
Pyrgo anomala
Quinqueloculina lamarckiana
Rotamorphina involuta 1
Sigmoilina distorta
Sigmoilina tenuis 1
Sigmoilopsis schlumbergeri 1 1 1 1 2 1 1 3 2
Siphonina reticulata 1 2 1 1 3 4 I
Sphaeroidina bulloides 1 10 5 3 4 2 2
Spiroloculina canaliculata 1 2
Textularia calva 1 I 1
Textularia conica
Textularia sagittula 1 1 1 1 1
Trifarina angulosa 1 1 2 1 1 3 I 1 1
Uvigerina auberiana 2 4 7 5 1 3 1 I 3 2 1
Uvigerina peregrina 15 5 11 4 7 7 15 10 6 8 23 1 5
Valvulineria bradyana 2 1 1 3 5
Miscellaneous 5 2 4 6 3 5 5 6 7 5 6 7 2 3 3 1
Total number counted 142 136 155 185 115 159 153 176 104 174 157 162 92 161 155 155 142 6.5

25
BREMER AND OTHERS

APPENDIX 2
Continued.

40 39 38 37 36 35 34 33 32 31 30 29 28 27 26 25 24 23

Ammonia beccarii 1 1
Asterigerinita mammila 1
Bigeneria nodosaria 1
Brizalina albatrossi 1 1 1 1 2 1
Brizalina attica 1 1 1
Brizalina catanensis 2 3 1 1 89
Brizalina difformis 1 4 I
Brizalina dilitata 1 2 3 7 1 2
Brizalina pseudoplicata
Brizalina alata 1 I
Bulimina aculeata + marginata 6 3 1 4 2 1 7 4 16 7 4 2 7 6 2 0
Bulimina costata 2 1 6 3 4 2 1 1 1 0 6 I 1 2 9 1 0 6
Bulimina exilis 1
Bulimina gibba 8
Cibicides boueanus 2 1 2 2 2 1 4 2 3 3 2 2 2
Cassidulina crassa
Cassidulina laevigata I1 20 8 15 4 2 10 14 10 7 7 5 13 9 3 4
Globocassidulina subglobosa 4 3 1 3 3 3 2 3 13 5 1 3 1 1 6 2 3
Cibicides lobatulus 1 4 2 2 1 1 1 1 1 1 1 4
Cibicidoides jloridanus 20 18 30 14 7 28 22 21 23 23 17 22 9 3 18 22 21
Eggerella bradyi 1 1 1
Elphidium crispum 1
Epistominella evax
Epistominella rugosa convexa 75 3
Florilus boueanum 1
Gavelinopsis praegeri 1 1
Globobulimina afJinis 2 1 1 1 5 1 5 2 2
Globobulimina pseudospinescens 1 3 1
Gyroidina altiformis
Gyroidina umbonata 3 4 2 2 1 4 1 1 4 10 1 1 1
Gyroidinoides neosoldanii 2 3 5 4 5 2 4 1 3 3 2 4 1 4 1 1
Hanzawaia rhodiensis 2
Hoglundina elegans 1
Hyalinea balthica 1 3 15 11 I 1
Amphicotyna scalaris 1 1
Melonis barleanus 6 3 1 1 3 1 3 2 1 1 2 8 3 1 1 6 3
Miliolinella circularis
Neoconorbina terquemi 1
Oridorsalis tener 1 3 1 1 1 1 3 2 1 1 2
Planorbulina mediterranensis 1
Planulina arimenensis 3 4 3 3 4 4 4 6 2 3 1 1 1 2 4 5
Pseudoclavulina crustata 1 3 I 1
Pullenia bulloides 3 3 1 2 4 1 1 6 1 2 1 1 2 1 2
Pullenia quinqueloba 1 1 1 1 1 1 1
Pyrgo anomala
Quinqueloculina lamarckiana 1
Rotamorphina involuta
Sigmoilina distorta 1
Sigmoilina tenuis I 1 1 1
Sigmoilopsis schlumbergeri 5 4 2 1 4 2 3 2 1 1 1 I 1
Siphonina reticulata I 1 2 1 4 1 1
Sphaeroidina bulloides 5 8 4 5 9 8 6 7 10 9 3 3 5 1 9 6 3
Spiroloculina canaliculata 1 1
Textularia calva 3 1 7 1 1 2 I 1 1
Textularia conica 1
Textularia sagittula 2 2 2 2 1 1 2
Trifarina angulosa 4 4 1 3 1 2 1 1 8 1 5 5 2 1 1 1
Uvigerina auberiana 2 4 5 2 1 1 5 1 5 6 2 1 4 2 4 1
Uvigerina peregrina 11 8 20 13 3 11 12 25 8 16 16 14 18 1 3 IO 7 10
Valvulineria bradyana 1 1 1
Miscellaneous 1 . 5 5 2 3 4 4 5 6 4 5 4 5 0 3 7 6 5
Total number counted 128 187 107 160 118 225 153 149 199 141 145 140 149 149 178 141 208 322

26
 PLIOCENE-PLEISTOCENE IN ITA1 .Y

APPENDIX 2
Continued.

60 59 58 57 56 55 54 52 51 50 49 47 46 45 44 43 42 4'
~

Ammonia beccarii
Asterigerinita mammila
Bigeneria nodosaria 3
Brizalina albatrossi 1 1
Brizalina attica 1
Brizalina catanensis 1 1 29 1 1
Brizalina dijjformis 1 1 1
Brizalina dilitata 1 20 5
Brizalina pseudoplicata
Brizalina alata 1 3 2 9 1
Bulimina aculeata + marginata 12 3 1 1 1 2 2 2 2 5 5 5 5 3 1 5 I 3 4 3 1
Bulimina costata 41 3 4 1 1 9 5 2 2 7 6 1
Bulimina exilis
Bulimina gibba
Cibicides boueanus 2 3 1 1 5 1 2 1 3 1 4
Cassidulina crassa 1 1 2 1 2 19 1
Cassidulina laevigata 2 6 10 9 11 27 7 14 15 14 10 10 4 2 15 11 20
Globocassidulina subglobosa 1 2 3 3 5 3 1 2 2 3 1 5 4 2 3 3
Cibicides lobatulus 1 1 1 1 1 3 2 2 1 1 1 3 1
Cibicidoides floridanus 6 17 10 16 8 14 10 14 3 4 4 7 20 24 18 14 23 49
Eggerella bradyi I 1
Elphidium crispum 1
Epistominella evax
Epistominella rugosa convexa
Florilus boueanum 2 2
Gavelinopsis praegeri 4 2 3 6 3 1 2 4 1
Globobulimina af$nis 1 2 1 2 4 3
Globobulimina pseudospinescens 1 2 1 2 1 1 1
Gyroidina altiformis 1
Gyroidina umbonata 1 1 1 1 2 1 1 1 2 1 2 2 4
Groidinoides neosoldanii 1 1 1 2 1 1 5 3 3 4 2
Hanzawaia rhodiensis 1
Hoglundina elegans 2 2
Hyalinea balthica 7 12 11 31 4 9 2 1 6 1 3 1 6 1 0 1 5
Amphicoryna scalaris I 1 10
Melonis barleanus 8 4 1 1 2 5 1 2 7 1 1 3 2 4 2
Miliolinella circularis 1
Neoconorbina terquemi 1 2 1
Oridorsalis tener 1 1 2 3
Planorbulina mediterranensis 1
Planulina arimenensis 4 7 7 6 1 3 2 4 2 1 1 5 3 6 9 3 2 1
Pseudoclavulina crustata 1 1 1 1 2 2 1 1
Pullenia bulloides 1 1 2 5 1 1 1 1
Pullenia quinqueloba 1 1 1 1 1 1 1 1 2
Pyrgo anomala 1 1
Quinqueloculina lamarckiana 3 2 1 1 1
Rotamorphina involuta
Sigmoilina distorta 3 1 2 2 1 1
Sigmoilina tenuis 2 2 1 1
Sigmoilopsis schlumbergeri 4 5 3 2 2 1 2 1 1 2 2 1 1 1 1
Siphonina reticulata 2
Sphaeroidina bulloides 8 7 3 4 5 6 5 2 1 6 1 4 2 7 5 3 2
Spiroloculina canaliculata 1 3 1 1
Textularia calva 4 4 1 1 1 2 1 4 3 1 1
Textularia conica 1
Textularia sagittula 1 1 1 2 3 2 2 1
Trifarina angulosa 1 5 6 8 5 1 5 1 3 2 4 4 2 8 8 7 9 1 0
Uvigerina auberiana 4 1 2 11 5 3
Uvigerina peregrina 5 12 14 13 10 14 25 12 13 1 12 28 15 4 16 9 1 7
Valvulineria bradyana 10
Miscellaneous 4 7 4 6 3 3 6 1 6 5 6 6 1 7 5 7 2 5
Total number counted 141 133 149 160 153 250 230 152 128 225 121 101 137 142 153 138 145 154

27
BREMER AND OTHERS

APPENDIX 3
Parker's (1958) present day eastern Mediterranean benthonic foraminiferal faunal census data as rearranged for this analysis (in three parts).
Values are percentages. Listed across the bottom are the percentages of the benthonic foraminifera in each of Parker's samples retained after
the species which do not occur at Le Castella were eliminated.

Sample 4647 4657 4718 4694 4723 4687 4685 4664 4660 4714 4656 4713 BS9 4654 4686 4702 4650 4670
Tax"" Denth in meters 366 384 541 561 561 631 647 658 665 665 680 684 731 746 199 799 859 996

Ammonia beccarii
Asterigerinita mammila
Bigeneria nodosaria 2 .3 4 2 1 .2 3 .5
Brizalina albatrossi .4 .4 1
Brizalina attica 1
Brizalina catanensis 5 2 1
Brizalina difformis .E
Brczalina dilitata 3 2 4 9 9 .5 6 3
Brizalina pseudoplicata .4 .2 5 .6
Brizalina alata
Bulimina aculeata + marginata 11 13 11 17 4 2 0 2 1 3 2 1 6 5
Bulimina costata 4 10 5 11 2 2 7 14 16 .4 1 .E .4 .5 .5 5 1
Bulimina exilis
Bulimina gibba 6 .6 1
Cibicides boueanus
Cassidulina crassa .7 2 5 2 2 1 . 5 4 1 7 4 1 2 5
Cassidulina laevigata 3 2 1 1 2 1 .5 .2 .2 .6
Globocassidulina subglobosa 1 2 1 2 1 2 .5
Cibicides lobatulus 3 1 1 . 4 2 1 1 .3 .2 2 1 .7 2 3
Cibicidoides floridanus 1 2 8 3 1 0 6 7 2 .3
Eggerella bradyi L
Elphidium crispum .L
Epistominella evax
Epistominella rugosa convexa 2 .2 2 8 13 3 2 3
Florilus boueanum
Gavelinopsb praegeri .7 .6 .7 2 .2 .2 .4 .7 .5 .5 .6
Globobulimina affinis
Globobulimina pseudospinescens 2 2
Gyroidina altiformis 8 3 4 . 7 2 2 3 9 7 2 1 5 3 5 3 2 4 3 2 6 5
Gyroidina umbonata .6 .7 .3 .3 .2 1 .7 1 .6
Gyroidinoides neosoldanii 3 5 2 4 . 4 6 2 3 1 4 5 6 1 1 4 5 2 1 3 2 0
Hanzawaia rhodiensis 2 .3 1 .2 3 1 .4 1
Hoglundina elegans 2 2 .2 .2 .7
Hyalinea balthica 2 2 . 4 1 3 1 2 . 6 .4 .5
Amphicoryna scalaris 5 2 2 . 7 2 . 6 2 .2 .6 1 .5 .6
Melonis barleanus 3 5 1 . 7 5 2 13 12 24 10 7 4 10 1 42 32 8
Miliolinella circularis .7 2 2 .E .6 1 2 1 1 2 2
Neoconorbina terquemi .2 .E .7 .5 2
Oridorsalis tener
Planorbulina mediterranensis .4
Planulina arimenensis 1 5 4 3 1 5 4 2 3 5 . 5 4 1 3 1 1 . 6
Pseudoclavulina crustata 5 1 3 . 8 2 1 2 1 . 2 1
Pullenia bulloides
Pullenia quinqueloba .6 2 1 .3 1 3 .6
Pyrgo anomala .6 2 .2 1 .2
Quinqueloculina lamarckiana 1 2 .3 .7 .3 .5
Rotamorphina involuta .4 .3 5 1 .7 .5 .3 1
Sigmoilina distorta 2 1 .4 2 .6 .2 1 .4 1 2 .4
Sigmoilina tenuis .3 3 .6 .6 1 .5 .6
Sigmoilopsis schlumbergeri 3 .3 .2 2 .6 .3 4 2 1 .6
Siphonina reticulata .7 .3 1 .4 .3 .3 .2 2 1 3 2 .6
Sphaeroidina bulloides 2 4 . 8 1 1 4 3 . 6 .3 4 1 2
Spiroloculina canaliculata .7 .4 1 1 .2 1 1
Textularia calva 2 .3 1
Textularia conica 2 . 3 1 . 2 1 1
Textularia sagittula 2 .4
Trifarina angulosa .7 1 4 .4
Uvigerina auberiana 3 5 9 3 .6 .6 3 . 5 7 1
Uvigerina peregrina 13 26 35 8 22 37 33 14 28 5 41 3 16 17 48 3 7 13
Valvulineria bradyana .3 .2 .5
Percent Accounted For 93 100 90 76 82 84 95 92 96 69 87 70 63 77 80 88 69

28
 PLIOCENE-PLEISTOCENE IN ITALY

APPENDIX 3
Continued.

Sample BS32 4662A 4666 4661 4658 4667 4657A 4649A 4654A 4668 4654B 4721A 4652 BS21A BS20 46578 BS29 4648
Taxon Depth in meters Bay 51 71 75 82 86 IC4 106 115 117 143 179 201 205 210 256 287 338

Ammonia beccarii 2 7 4 2 5 .5 2
Asterigerinita mammila 4 13 4 38 5 2 3 .5 16 11 9 4
Bigeneria nodosaria .6 .6 2 2 .2 6 s 1 1 2 5 3 . 7
Brizalina albatrossi
Brizalina attica .2 I .5 .7 .6 2 .4
Brizalina catanensis 5 4 8 3 8 .3 2 2 8 4 2 2
Brizalina difformis .5 I 3 .3
Brizalina dilitata 2 1 2 3 1 1 3
Brizalina pseudoplicata .3 .3 .5
Brizalina alata
Bulimina aculeata + marginata .2 .6 6 . 3 2 2 6 5 3 7 5 7
Bulimina costata .6 .6 1 .6 1 3 4 9 1 0 1 5
Bulimina exilis
Bulimina gibba 1 4 2 4 1 1 4 3 7 3 9 5 1
Cibicides boueanus
Cassidulina crassa 3 .5 2 3 3 .3 .7 I .4
Cassidulina laevigata 1 1 0 1 7 7 8 5 1 14 5 3 2 1 4 I
Globocassidulina subglobosa 1 .4
Cibicides lobatulus 10 3 7 6 4 . 6 7 1 4 3 1 5 5 9 1 1 3 2 2
Cibicidoides floridanus .7 .5 I .3 .4 1 4 3
Eggerella bradyi
Elphidium crispum 7 1 4 2 3 2 2 .4 5 6 .2 .4
Epistominella evax
Epistominella rugosa convexa
Florilus boueanum
Gavelinopsis praegeri .3 .3 2 3 1 2 5 2 .3 I .3 I
Globobulimina affinis
Globobulimina pseudospinescens
Gyroidina altiformis . 7 . 6 2 1 2 1
Gyroidina umbonata .3 1 . 2 2 1
Gyroidinoides neosoldanii .3 .3 .7 2 2 1 3
Hanzawaia rhodiensis 6 7 2 7 5 5 4 1 .5 1 .7 .7 .3 .3 .7
Hoglundina elegans .2 1 .3 .5 I .7 2 .3 .4
Hyalinea balthica 2 2 3 6 3 6 2 6
Amphicoryna scalaris .3 .3 1 .5 . 3 3 2 1 1 4 2 . 4
Melonis barleanus 4 8 1 6 5 5 3 3 1 5 1 1 2 1 2 7 8 7
Miliolinella circularis 2 .6 .3 1 .6 .3 .3 1 4 .4 2 .5 3 2 2 .6 2 .4
Neoconorbina terquemi 2 7 5 3 4 5 1 1 3 .3 1 .6 .3 .3 1
Oridorsalis tener
Planorbulina mediterranensis 13 1 2 . 2 2 4 I .7 .4 .6 .3 .3 .7 .6
Planulina arimenensis .3 2 1 . 4 2 . 5 3
Pseudoclavulina crustata .5 2 . 3 . 7 I 2 I
Pullenia bulloides
Pullenia quinqueloba .2 .5 .3 .3 .3 .4
Pyrgo anomala 1 1 1 3 .5 .6 .7 .2 .3 .5 1
Quinqueloculina lamarckiana .6 .5 1 .5 1 .3 .6 .3 .2 2 2 I
Rotamorphina involuta
Sigmoilina distorta .2 I .2 .6 7 .5 .3 .3 1 I 1
Sigmoilina tenuis .3 .4
Sigmoilopsis schlumbergeri 2 .6 .7 .2 I 1 1
Siphonina reticulata 1 .7 .5 1 .2 1 1 .4
Sphaeroidina bulloides .2 .3 2 . 5 2 4 2 3 3
Spiroloculina canaliculata .3 .6 I .6 .5 .3 .4 .3 .7 .3 .5
Textularia calva 2 3 4 3 3 2 2 5 3 2 5 2 5 2 2 2 2 2
Textularia conica .4 1 2 .3 .3 .3 2 .4 3 2 .5 1
Textularia sagittula 1 .3 7 4 2 5 2 2 9 6 5 2 2 . 3 2 . 4
Trifarina angulosa 2 .6 .4
Uvigerina auberiana 1 2 1 3
Uvigerina peregrina .6 .3 3 IO .5 5 11 21 22 19 9 16
Valvulineria bradyana .2 2 .3 1 .3 .4 2 2 .3 .3
Percent Accounted For 37 60 53 72 63 64 67 78 46 43 61 65 87 87 88 95 83 89

29
BREMER AND OTHERS

APPENDIX 3

Continued.

Sample 4671 4693 4649 4688 4716 4705 4673 4678 4683 4689 4698 4701 4681 4710 4711 4708 4676
Taxon Depth in meters 1016 1073 1265 1312 1378 1788 1844 1862 2358 2442 2738 2760 2852 3241 3309 3499 3974

Ammonia beccarii
Asterigerinita mammila
Bigeneria nodosaria
Brizalina albatrossi 4 2
Brizalina attica
Brizalina catanensis .3
Brizalina difformis
Brizalina dilitata 2 2
Brizalina pseudoplicata 50
Brizalina alata
Bulimina aeuleata + marginata
Bulimina costata 2 6 3
Bulimina exilis
Bulimina gibba
Cibicides boueanus
Cassidulina erassa .6 2
Cassidulina laevigata
Globocassidulina subglobosa .3
Cibicides lobatulus .6 4 2
Cibicidoides jforidanus 4 2 8
Eggerella bradyi
-Elphidium crispum
Epistominella evax
Epistominella rugosa eonvexa .6 3
Florilus boueanum
Gavelinopsis praegeri .6
Globobulimina affinis
Globobulimina pseudospinescens
Gyroidina altiformis 5 4 34 11 12 18 14 23 5 8
Gyroidina umbonata .3
Gyroidinoides neosoldanii 9 36 15 67 49 82 64 19 5 18 6
Hanzawaia rhodiensis 1 2
Hoglundina elegans 16 .6
Hyalinea balthica
Amphicoryna scalaris
Melonis barleanus 5 14 6 13
MiliolineJla circularis 3 4 2 5
Neoconorbina terquemi
Oridorsalis tener
Plan orbulina mediterranensis
Pia nul ina arimenensis 4
Pseudoc1avulina crustata
Pullenia bul/oides
Pullenia quinqueloba .6
Pyrgo anomala
Quinqueloculina lamarekiana
Rotamorphina involuta
Sigmoilina distorta 45
Sigmoilina tenuis .3
Sigmoilopsis schlumbergeri .6
Siphonina reticulala 1
Sphaeroidina bulloides 1 2 .6
Spiroloeulina canaliculata .3
Textularia calva
Textularia eonica
Textularia sagittula
Trifarina anulosa
Uvigerina auberiana .6
Uvigerina peregrina 23 7 3 2 25 6 6
Valvulineria bradyana
Percent Accounted For 82 54 80 78 73 100 84 42 10 59 18 50 0 12 50 0 13

30