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Taxomorph: Foraminifera from the South-west Pacific. An inter-active

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 TAXOMORPH

FORAMINIFERA
FROM THE
SOUTH-WEST
PACIFIC

An introduction to the Foraminifera

Kingdom Protozoa
Phylum Foraminifera
 2

CONTENT PAGE

Introduction………………………….…………...…..……….. 3
The living foraminifer…………………………………………. 5
Life history of foraminifera…………….………...…………… 7
The test……………………………….………………….…. 10
Wall structure…………………………………….…….. 11
Chitinous wall………………..…………….……..12
Agglutinate wall…………………………….…….11
Microgranular calcareous wall……………..……. 12
Calcareous porcelaneous wall…………………….13
Calcareous hyaline wall………………………..…13
Mode of Growth (Chamber arrangement)…………...… 14
Unilocular……………………………………..…. 14
Multilocular………………………………...……. 14
Uniserial………………………………….…..….. 15
Biserial…………………………………..…..……15
Triserial……………………………..………..….. 15
Planispiral………………………………….…..… 15
Trochospiral……………………………….…….. 16
Biloculine………………………………………... 17
Triloculine……………………………………..… 17
Quinqueloculine…………………………….…… 18
Spiroloculine……………………………….……. 18
Ornamentation……………………………………..….. 19
Aperture………………………………………….…….. 20
Method of study……………………………………………… 22
Living assemblages…………………………………….. 23
Benthic foraminifera……………………..….…... 23
Pelagic foraminifera…………………..…………. 24
Total assemblages……………………………..…..…… 25
Washing……………………………………….…. 25
Separation…………………………………….…. 26
Selecting and mounting………………….……..…26
Statistical studies…………………………..…………... 28
Classification……………………………………………29
Further readings…………………………………………………….…….. 30
Appendix - Geologic time scale…………………………………….……. 36
 3

INTRODUCTION

The Foraminifera are unicellular organisms living predominantly in the marine

environment. Their size ranges from 0.05 to 2 millimeters and only a few species
reach greater dimensions. Most foraminifera build a test (shell) of mineral matter
(predominantly calcium carbonate) and these tests are commonly preserved as
fossils in sedimentary rocks.

The first record of foraminifera goes back to the 15th century BC when Herodotus
noted their presence in the blocks of rock forming the Egyptian pyramids, but, as
with all other fossils, it is only much later that they were generally recognized as
the remains of living organisms. With the invention of the microscope (1660) a
new field of observation and study was opened and foraminifera were among the
organisms studied and described by early naturalists although classified as
cephalopods, corals or worms. To this group of authors belong Linnè, Lamarck
and d'Orbigny, the latter being the proposer of the name Foraminifera for these
organisms.
Since then, various aspects of foraminifera have been extensively studied and
their usefulness in stratigraphy was gradually realised; they have a place in
Applied Science as stratigraphical index fossils, widely used in petroleum
exploration. Their usefulness is due to their abundance in the marine sedimentary
rocks, with which petroleum deposits are usually associated, and to their small
size, which enables representative samples of a fauna to be obtained from a small
sample of rock. This is a feature of great value in oil field prospecting and
development when only small cores or chips of rock from the drilling are
available.

The importance of foraminifera as index fossils is related to restricted time-range

occurrence of many species in the geological history. Their importance is
increased if, at the same time, they were geographically widespread.
Other species, characteristic of a particular environment, may be used as guide
fossils in palaeogeographical reconstruction.
 4

In recent years palaeontologists and zoologists have realized the need for a better
understanding of the ecology of living foraminifera, instead of their geologic
occurrence only.
Foraminifera are also important in paleoclimate, paleosealevel, paleodepth and
paleoceanography studies and, recently, in absolute age determinations using
oxygen isotopes.
Statistical studies, based on the distribution of benthic foraminifera in the marine
and estuarine environments, have also shown that these organisms can be
successfully used to identify the various ecological provinces, to detect
environmental stress conditions and to monitor pollution.

Foraminifera occur in nearly all the aqueous environments, mainly in marine and
in brackish waters. Their abundance varies considerably and may reach more than
2,500 living individuals per square centimetre of sea bottom.

Because their existence is related to many factors, such as oxygen supply,

temperature, salinity, food supply, depth of water, each environment shows a
definite foraminiferal fauna. These factors are related to the characteristics and the
circulation of the various water masses, which therefore are reflected in the
distribution of foraminiferal assemblages.
 5

THE LIVING FORAMINIFER

The soft body of a foraminifer consists of a mass of protoplasm whose cytoplasm

is differentiated into an inner dark-coloured endoplasm and an outer layer of clear
ectoplasm, which is restricted around the aperture and forming a thin layer outside
the test. One or more nuclei may be present but these are always located within
the test. From the ectoplasm many projections extend outwards forming
sometimes a fine reticulate pattern (anastomosing pseudopodia) (fig. 1).
The pseudopodia are mainly used for the capture of food, respiration, and the
expulsion of waste products as well as for moving around or as attachment to the
substratum.

Fig. 1 – An unilocular (single chambered) foraminifer.

The test, which will be described later, is secreted by the ectoplasm and may vary
considerably in shape. It may consist of only one chamber (unilocular tests) or
many chambers (multilocular tests).
In the multilocular tests a rounded chamber is initially formed (proloculus);
subsequent additional chambers are formed according to a pattern of growth
which is characteristic for each species.

The foraminifera feed generally on diatoms, larvae of copepods, algae, infusoria,

flagellates and bacteria that are captured by the pseudopodia; digestion may occur
either outside or inside the test. In some shallow water foraminifera the presence
 6

of symbiotic algae is also noted. The majority of foraminifera live on the sea floor
at all depths (benthic foraminifera), some moving very slowly by means of the
pseudopodia (few millimetres per hour) others attaching themselves to the bottom
or to other organisms (seaweed, corals, molluscs). Other species live in the
surface waters (planktonic foraminifera) and are transported by the ocean
currents.
 7

LIFE HISTORY OF FORAMINIFERA

The life history of a few species of foraminifera are known, although many
thousand of living species have been described from the test alone.
The initial investigations of the life history were made independently by two
palaeontologists De La Harpe and Hantken. They noted the constant presence in
Nummulites of two different forms; one with a small proloculus and large test
(microspheric form) and another one with a large proloculus and relatively
smaller test (megalosphaeric form). These came to be recognized later as
dimorphic forms of the same species. They generally represent phases of
alternation of generations.
A generalised life-cycle is shown in figure 2.

Figure 2 – A generalised life-cycle

 8

Let us start from a microspheric individual (1) with a small proloculus and a large
test in its adult stage. At the time of reproduction the nucleus and cytoplasm
divide, a portion of the cytoplasm surrounding each nucleus (2). Each young (3),
so formed, begins to form a new test (4). This first part of the reproduction cycle
is assexual. The young so produced are relatively large and they will secrete a
large initial chamber, which will become the proloculus of megalospheric
individuals (5). New chambers will be added to this proloculus to eventually form
a megalospaheric adult (6). When the megalospheric individual reproduces the
nucleus and cytoplasm divide into a number of gametes, which are free living,
flagellate and lack a test (7). This phase of the reproduction is sexual and the
fusion of two gametes (8, 9) from two individuals produces a small zygote (10).
This zygote secretes a test (11), which will be the proloculus of a new individual
of the microspheric generation (12).
Recent studies have also shown the existence, in at least some species, of a
trimorphic cycle with two assexual phases and with the formation of three
individuals, two megalospheric individuals and one microspheric.

Fig. 3 – Plastogamic pair Fig. 4 – Float chamber

Variations occur in the mode of reproduction; in some foraminifera two

individuals form a pair (plastogamic pair) attaching themselves together by the
ventral side, before exchanging the gametes. By so doing, they form a protected
 9

area between them where the conjugation of the gametes occurs (Fig.3). In others
the discharge
of the gametes occur in a pelagic stage. To achieve this condition the foraminifer,
at the time of reproduction, forms on its ventral side a large clear chamber which,
filled with gas, allows the animal to float. The gametes are discharged into the
float chamber and eventually leave it through the large pores on its wall (Fig.4).
 10

THE TEST

The test is a hard protective shell that is secreted by the animal and because it is
the only hard part of the organism it is the only remain found fossil.
Vast numbers of tests of dead foraminifera accumulate in the sea floor sediments
and are often studiedat this stage prior to their deep burial and fossilisation in
rocks. During these processes many tests are destroyed so that the pre-fossil
abundance is generally much greater than the fossil assemblage.
The first studies on foraminifera were done by palaeontologists who therefore
developed a classification based on the shape of the test. Recent higher
classifications of the foraminifera based on biological studies are in general
agreement with the classification based on the morphology of the test. For all
purposes, therefore, the test is the most important taxonomic feature and its
detailed structure and morphology are outlined.
The descriptive terms are used, in the catalogue, to help in the identification
process.
 11

Wall Structure
Five distinct main types of shell wall occur in foraminifera.

Agglutinate wall. This wall is constructed with sand grains (arenaceous), or other
particles, usually cemented by calcium carbonate (Fig. 5).
The extraneous particles may be fine sand grains (usually quartz), heavy minerals,
diatoms, fragments of molluscan shells, sponge spicules and many other types of
debris, even foraminiferal tests. Some foraminifera select particular types of
material from the range available in the environment. A few foraminifera found in
the Arctic Ocean utilize only mineral grains transported by icebergs, some utilize
grains of equal dimensions and others use grains of the same colour. In the same
bottom conditions, for example, Rhabdammina uses sand grains while Marsipella
only sponge spicules and Psammosphera mostly mica flakes.

Fig. 5 – An agglutinated (arenaceous) test

In some foraminifera selection does not appear to be vital. A relatively large

opening, called the aperture, forms the only link between the interior and the
exterior of the test.
 12

Chitinous wall. The chitinous wall, thin and easily deformable, is present in the
simplest forms and in the juvenile stage of few more complex ones. The
occurrence of foraminifera with this type of wall in the fossil record is not
common and therefore this wall type is not considered in the catalogue

Microgranular calcareous wall. This particular type of test consists of very

small irregular grains of calcite closely packed and cemented together by calcium
carbonate. It is typical of Palaeozoic calcareous foraminifera.
 13

Calcareous porcelaneous wall. This type of wall is formed by small crystals of

calcite with a random orientation but generally parallel to the surface of the test,
which has an opaque porcelaneous appearance (Fig. 6).
As in the agglutinate forms, the only opening is the aperture.
Calcareous hyaline wall. In this case, the crystals of calcite forming the test are
all orientated with the long axis at right angles to the surface of the test (Fig. 7).
This type of wall shows numerous perforations (pores) of very small diameter so
that is also termed "calcareous, perforate" while the calcareous porcelaneous test
is defined as "calcareous, imperforate". These pores, together with the aperture,
function as passages for the cytoplasm carrying food or waste products between

Fig. 6 – A calcareous porcelaneous test Fig. 7 – A calcareous hyaline test

the interior and the exterior of the test. In the imperforate tests such protoplasmic
movement occurs only through the aperture.
In the case of calcareous hyaline tests, when a new chamber is formed, a thin
layer of shell material is added to the entire earlier part of the test, whereas in
foraminifera with wall structure of the other types, this enclosing layer is not
formed. It results in the walls of the early chambers of calcareous hyaline
foraminifera being much thicker than the walls of later chambers.
 14

Mode of Growth (Chamber arrangement)

The arrangement, shape and number of the chambers may vary greatly but each
genus has its own characteristic pattern. Most of the terminology used is
descriptive and uses common words. However, a few terms specifically used for
this group of organisms require some explanations.

The test may be composed by one chamber (unilocular) (Fig. 8) or two or more
chambers (multilocular). The main opening of the test is called the aperture.

In case of multilocular tests the chambers may be arranged in a variety of ways.

When each new chamber is added, part of the external wall of the previous
chamber becomes an internal partition (septum) between the two adjacent
chambers. The linear junction between two adjoining chambers is termed a suture;
spiral and radial sutures may occur (Fig. 9).

Fig. 8 – An unilocular test Fig. 9 – Various sutures

The previous aperture becomes an internal opening between chambers called the
foramen, from which the name foraminifera is derived.
 15

Among the most common patterns of growth the following are noted:
Uniserial, with chambers added in line which may be straight (Fig. 10A) or
curved.
Biserial, with chambers added in rows of two (Fig. 10B). In some cases the
biserial test may become uniserial in adult stage.

Fig. 10 – (A) uniserial, (B) biserial tests Fig. 11 – Triserial test

Tests may be triserial, with three chambers to a whorl (Fig. 11).

The chambers may also be added in a coil tangent to a plane (planispiral) or to a
cone (trochospiral). The planispirally coiled test may become uncoiled in adult

Fig. 12–Test planispiral becoming uncoiled Fig. 13 – Septal bridges.

 16

stage (Fig. 12). In other planispirally coiled forms, such as Elphidium, each
chamber projects across the preceding suture several extensions which appear as
small ridges (septal bridges) over the suture itself (Fig.13).
These sutural extensions may end blindly against the wall of the preceding
chamber (septal bars) or they may open into the earlier chamber by means of a
small pore (retral processes).

In trochospiral tests growth occurs coiling around a cone; the side which shows
all the previous whorls (evolute) is called the dorsal side (Fig. 14) while the one
showing only the chambers of the last whorl (involute) is called the ventral side
(Fig. 15). The portion of chamber in which the aperture opens is the apertural
face.

Fig 14 – Dorsal (evolute) side Fig. 15 – Ventral (involute) side

The ratio between diameter of the cone and its height may be used to define the
type of trochospiral growth; in high trochospiral the cone has a small apical angle,
and the ratio height / diameter greater than 1 (Fig. 16); while in the low
trochospiral the apical angle is large with the ratio height / diameter smaller than 1
(Fig. 17).
 17

Fig. 16 – High trochospiral test Fig. 17 – Low trochospiral test

A group (miliolids) shows another mode of growth. In the milioline arrangement,

the chambers are added using a number of planes at different angles :
biloculine = planes are 180° apart, only the final two chambers are visible (Fig.
18);
triloculine = planes are 120° apart, three visible chambers (Fig. 19);

Fig. 18 – Biloculine test Fig. 19 – Triloculine test

 18

Fig. 20 – Quinqueloculine test Fig. 21 – Spiroloculine test

quinqueloculine = planes at 144°, five visible chambers ( Fig. 20);

spiroloculine = initially milioline followed by chambers added at opposite sides
of a single plane (Fig. 21).
 19

Ornamentation
The ornamentation of the test may consist of spines (Fig. 22), costae or ribs (Fig.
23), papillae (Fig.24) and keels (Fig. 25), all produced by a thickening of shell
material.

Fig. 22 – Spines Fig. 23 – Costae or ribs

Fig. 24 – Papillae Fig. 25 - Keel

Sometimes, in specific determinations, it is necessary to consider the alternation

of generations. As mentioned earlier, this may cause the appearance of two or
more forms with slightly different tests.
 20

The Aperture

The aperture is another important feature from morphological and phylogenetic

points of view. The most common types are simply at the end of the final chamber
(terminal) (Fig.26) or at the base of the apertural face either as a single long slit
(Fig. 16) or as a series of large pores (Fig. 27) or cribrate (Fig. 28).

Fig. 26 – Terminal aperture Fig. 27 – Pores (at base of apertural face)

Fig. 28 – Cribrate aperture Fig. 29 – Aperture with neck

 21

Sometimes the aperture is at the end of a neck (Fig. 29) or with an entosolenian
tube, which consists of an internal tube departing from the aperture.
Often the aperture shows the presence of an extension called a tooth (Fig. 19),
which may be simple, bifurcate or plate like.
 22

METHOD OF STUDY

The Commonwealth of Australia comprises six States and two mainland Territories.
Under the Constitution, control of Australian flora and fauna rests with the States and
Territories. The collecting and moving of plants and animals in Australia is managed
by State, Territory or Commonwealth licensing authorities in each jurisdiction. Before
collecting, it is essential to obtain the permission of each relevant authority.
However, due to the nature of the sample used for studies of recent foraminifera,
collecting outside National Parks may not require special permission.

ABRS provides a list of contacts for obtaining permits for collecting flora and fauna.
It is available from the ABRS website
http://www.ea.gov.au/biodiversity/abrs/about/publications/guide/index.html.
Environment Australia manages permits for collecting in Commonwealth national
parks regulated by the Environment Protection & Biodiversity Conservation Act
1999, and details can be obtained from
http://www.ea.gov.au/epbc/permits/index.html.

In New Zealand, no collection is allowed in a Marine Reserve without obtaining

a written permit from the relevant regional office of teh Department of
Conservation. See http://www.doc.govt.nz/cons/marine/reserv.htm

The method of collecting, preparing and studying recent foraminifera varies

according to the nature of the material and the purpose of the investigation.
Studies of foraminifera are generally directed either towards an understanding of
the seasonal variations of populations in a selected environment or towards
defining their ecological or biogeographic distribution patterns.

In the first case the object of study is the population living at the time of collecting
and therefore it is necessary to preserve the soft protoplasm so as to be able to
discriminate living from dead specimens. In the second type of research the total
population is considered and therefore only the test is necessary.
Irrespective of the type of study, species identification is carried out on the
morphology of the test.
 23

Living assemblages

Benthic foraminifera

Several methods of collecting are used and they vary according to the type of area
to be sampled.
Samples could be collected by gently pushing a coring tube, such as a short
length of plastic piping, into the sediment to a depth of 10-20 cm taking extreme
care not to disturb the natural position of the surface of the sediment inside the
tube. A tube of 5 cm in diameter is adequate.
In deeper waters, samples may be collected by means of a dredge, grab or corer,
but there is great likelihood of disturbing the surface layer of the sediment.

As soon as possible after collection, the sample should be placed in a small plastic
bag, or small glass jar with sealable top, after removing as much seawater as
possible. A solution of alcohol (70%) and Rose Bengal stain (0.1%) is then added
in order to preserve and stain the living foraminifera. The protoplasm of any
living foraminifera will be stained a deep pink colour. It is advisable to put into
the plastic bag, containing the sample, an amount of solution at least double the
volume of the sediment and to mix it together by gentle shaking. The bag is then
sealed and may be stored.
Examination of the stained foraminifera can be made any time after half an hour
has elapsed. It is recommended rinsing out the stain from the sample and, if the
sample is muddy, washing out the mud with water just prior to examination in a
shallow dish with water.
The sample is examined wet under a stereomicroscope and the foraminifera,
revealed by the staining to have been alive at the time of collection, are removed
with a fine brush or by finger operated suction on a glass capillary tube and
mounted in slides.

As algae or bacteria sometimes line empty tests, the staining method using Rose
Bengal may produce erroneous results as these tests become stained and therefore
are considered as alive at the time of collection.
 24

A greater accuracy is obtained by using, as the staining agent, a saturated solution

of Sudan Black B in 70 percent ethanol. The use of this stain, however, requires
temperature control and therefore is generally carried out under laboratory
conditions. A fixative is needed to preserve the foraminiferal protoplasm during
the period between collection and preparation; a basic fixative must be used as
decalcification occurs if the tests are exposed to an acid solution (pH<4, even
pH<6).

Pelagic foraminifera

The collection is done with a fine plankton net, which is towed behind a vessel for
a certain length of time. The catch will consist of copepods, fish larvae and many
other planktonic forms besides foraminifera.

Usually several workers study a catch of plankton so any methods leading to the
destruction of all organisms except those to be studied are not desirable.

A useful method of concentrating foraminifera is to use a saturated solution of

NaCl. The sample is washed in water to remove the buffered formalin in which it
is normally preserved, and then it is placed in a large beaker containing the
saturated solution already prepared. The foraminifera are among those organisms
which sink, being so separated from the catch. The floating part is then placed
again in buffered formalin or ethanol and preserved for further study.
 25

Total assemblages

In this case the preservation of the organic matter is not important and the material
may be easily collected without special equipment. Along the beaches a natural
concentration of foraminifera, although often partially broken, may be found in
the whitish swash line left by the receding waves. In shallow waters a small
quantity of bottom sediment is easily obtainable with a small dredge.

The material so collected may be then washed with fresh water, dried in a warm
oven and then stored without any deterioration.

Washing

To examine the foraminiferal content it is necessary to wash the sample carefully

in order to remove clay and organic matter. To clean foraminifera from Recent
sediment is generally sufficient to wash the sample through a sieve (64 micron to
the inch) with the aid of a gentle spray of water to remove all mud.
Sometimes it may be necessary, to obtain clean tests, to boil the sample very
gently in water with a small amount of washing soda (Na2CO3). A few minutes
boiling may be sufficient.
Samples containing fossil foraminifera may need to be boiled for several hours; it
is recommended boiling in H2O2 or washing soda to disaggregate fossil samples
of medium hardness. For cemented sedimentary rocks the study is generally
carried out using thin sections.

For recent sediments, the sand fraction is then dried and it is often subdivided in
different size fractions through a set of sieves of 40, 80 and 150 meshes to the
inch (this can be done under either dry or wet conditions).
The reason for subdividing the sample in three size ranges is because the
examination, identification and counting of the various individuals, using a
stereomicroscope, can be carried out more easily if the tests under view vary in
size within a limited range.
 26

Special care should be taken in thoroughly cleaning the sieves after each use in
order to avoid any contamination. It is recommended dipping wet sieves in a
solution of methylene blue between sample preparation to blue stain any shells
from previous samples or any shell contaminants.

Separation

The foraminiferal tests may be concentrated in different ways. A very useful

method especially for Recent sediments, is the floating method. Each dry fraction
of the sample is gently poured in a beaker containing a heavy of liquids, the most
common but also the most dangerous used to be carbon tetrachloride (as an
ozone-depleting chemical it is now being phased out); the floating fraction,
containing the foraminiferal tests present in the sample, is easily decanted into a
filter paper and dried. All such work should be done under the most strict
laboratory conditions and in an approved fume cupboard as the vapours of CCl4,
are highly poisonous.
Rubber gloves must be used at all times as the carbon tetrachloride can be
absorbed through the skin. The floated material containing the foraminifera is
collected on a filter paper and left to dry (still in the fume cupboard). Once fully
dry, the concentrate can be transferred into small jars and it is ready for study
under a binocular microscope.
An alternative, but more expensive, heavy liquid sometimes used is sodium
polytungstate.

Selecting and Mounting

The sediment containing the foraminiferal tests is placed in a picking tray and is
examined under a stereomicroscope at 10-20 times magnification. The
foraminiferal tests may be picked up with a very fine moistened brush (000 size)
and mounted on cardboard or plastic slides on which the tests are generally glued
with gum tragacanth (a water soluble natural gum).
Cardboard or plastic slides with black mounting backing and white printed grid
are available from a number of suppliers:
 27

Australia - The Biology Shop, 15 Mount View Ave., Hazelbrook, NSW 2779;
phone (02) 4758 6765, fax (02) 4758 6806 (plastic slides).
New Zealand - Rice Printers, Rostrevor St, Hamilton, Phone 07 838 1229
(cardboard slides).
A glass cover is generally placed on the slide (Fig. 30), using a metal holder in
case of cardboard slides, or by sliding a glass microscope slide into the space
provided in case of plastic slides.

1 2 3 4 5 6 7 8 9 10

11 12 13 14 15 16 17 18 19 20

21 22 23 24 25 26 27 28 29 30

31 32 33 34 35 36 37 38 39 40

Fig. 30 – Microfossil slide.

Gum tragacanth is sold in a powder form and can be dissolved in warm water; a
few drops of alcohol or any other anti-bacteria liquid should be added prior its
use. The gum solution is generally brushed on in several coats of the slide and
allow to dry before use. The foraminiferal tests are then transferred to the slide in
a wet state, remaining glued to the slide itself.
 28

Statistical studies

Detailed sampling is necessary for a proper numerical treatment of the data. The
method most commonly used is sampling an area according to a grid pattern or
along transects; the spacing depends upon the details to be obtained and the size
of the area.

To make possible a direct numerical comparison between stations a constant

volume of sediment (10-40 cm3) should be processed from each station. About
300 individuals are regarded as representative of the station, although this value
may vary according to location. Generally the volume of sediment considered
contains a much larger assemblage which is then split. If, however, the number of
individuals is not sufficient, an additional volume of sediment is processed.

All the individuals must be identified and counted and the total population of each
species expressed as percentage value of the total assemblage of the station. These
percentages form the initial data base for the numerical treatment.
 29

Classification

Foraminifera are largely identified and classified on the basis of their shell
morphology.
The early foraminiferal literature from Agricola (1558) and Beccarius (1731)
regarded these organisms as worms, cephalopods, gastropods or even corals.
The earliest classification was that of De Blainville (1825). Some of the
classifications that followed were those of d’Orbigny (1826) [who used the term
foraminiferés for the first time], Schultze (1854), Reuss (1862), Schwager (1877),
Brady (1884), Cushman (1925), Chapman and Parr (1936), Glaessner (1945),
Hofker (1951) and Loeblich and Tappan (1964).

The classification adopted in this catalogue is that of Loeblich and Tappan (1988)
for genera and families and Loeblich and Tappan (1992) for orders.
 30

Further readings

For a complete and in depth understanding of this group of organisms a number of

textbooks and reference papers are suggested.

Textbooks
Loeblich, A. R. and Tappan, H. 1964. Sarcodina chiefly "Thecamoebians" and
Foraminiferida. Treatise on Invertebrate Paleontology, Protista. R. C. Moore.
Lawrence, Geological Society of America and University of Kansas Press.
Loeblich, A. R. and Tappan, H. 1988. Foraminiferal Genera and their Classification.
New York, Van Nostrand Reinhold Company.
Murray, J.W. 1973: “Distribution and ecology of living benthic foraminiferids.”
Heinemann, London, 288 pp.
Murray, J.W. 1991: “Ecology and paleoecology of benthic foraminifera”. Longman
Scientific and Technical, Avon, 397 pp.
Boltovskoy, E.; Wright, R.C. 1976: “Recent foraminifera.” Junk, The Hague, 515 pp.
Sen Gupta, K.B.(Ed) (1999). Modern Foraminifera. Kluwer Academic Publisher,
Dordrecht, The Netherlands.

Classification
Brady, H.B. 1884. Report on the foraminifera dredged by H.M.S. Challenger during
the years 1873-1876. Challenger Expedition, Reports, Zoology. 9.
Chapman, F. and Parr, W.J. 1936. A classification of the Foraminifera. Proceedings
of the Royal Society of Victoria, 49 (1): 139-151.
Cushman, J.A. 1925. An introduction to the morphology and classification of the
Foraminifera. Smithsonian Miscellaneous Collection, 77 (4): 1-77.
De Blainville, H.M. 1825. Manuel de malacologie et de conchyliologie; 664 p., F.G.
Levrault (Paris).
d'Orbigny, A. 1826. Tableau methodique de la classe des Cephalopodes. Annales des
Sciences Naturelles, 7 (1): 245-314.
Glaessner, M.F. 1945. Principles of micropaleontology. Melbourne University Press:
pp.296.
Hofker, J. 1951. The Foraminifera of the Siboga Expedition. Part III. Siboga-
Expeditie, (Monographie 4a): 1-513.
 31

Loeblich, A.R. and Tappan, H. 1992 - Present status of foraminiferal classification.

In: Takayanagi, Y. and Saito, T. (eds) Studies in Benthic foraminifera.
Proceedings of the Fourth Symposium on Benthic Foraminifera, Sendai, 1990.
Tokai University Press: pp. 93-102.
Reuss, A.E. 1851. Ueber die fossilen Foraminiferen und Entomostraceen der
Septarienthone der Umgegend von Berlin. Zeitschrift der Deutschen
Geologischen Gesellchaft, Berlin, 3: 49-91.
Schultze, M.S. 1854. Ueber den Organismus der Polythalamien (Foraminiferen),
nebst Bemerkungen über die Rizopoden im Allgemeinen. Engelmann, W.:
Leipzig.
Schwager, C. 1877. Quadro del proposto sistema de classificazione dei foraminiferi
con guscio. Bollettino del Regio Comitato di Geologia, Italia, 8 (1-2): 18-27.

Reference papers (restricted to the South Pacific area)

Studies on ecological distribution of Recent foraminifera of brackish or sheltered

harbour environments
Albani, A.D. 1968: Recent Foraminifera from Port Hacking, New South Wales.
Contributions from the Cushman Foundation for foraminiferal research 19(3):
85-119.
Albani, A.D. 1978: Recent foraminifera of an estuarine environment in Broken Bay,
New South Wales. Australian journal of marine and freshwater research 29:
355-398.
Albani, A.D., 1993 - Use of benthonic foraminifera to evaluate the environmenta
characteristics of Port Hacking: significance for coastal engineering works and
pollution; pp. 28-37. In McNALLY, G., KNIGHT, M. and SMITH, R, (Ed),
Collected Case Studies in Engineering Geology, Hydrogeology and
Environmental Geology, Geological Society of Australia , 137 pp.
Hayward, B.W.; Grenfell, H.R.; Cairns, G., Smith, A. 1996: Environmental controls on
benthic foraminiferal and thecamoebian associations in a tidal New Zealand inlet.
Journal of foraminiferal research 26(2): 150-171.
Hayward, B.W.; Triggs, C.M. 1994: Computer analysis of benthic foraminiferal
associations in a tidal New Zealand inlet. Journal of micropalaeontology 13: 103-
117.
 32

Studies on ecological distribution of Recent foraminifera of deep-sea environments

Hayward, B.W., Carter, R., Grenfell, H.R., Hayward, J.J., in press. Depth distribution
of Recent deep-sea benthic foraminifera east of New Zealand, and their
potential for improving paleobathymetric assessments of Neogene microfaunas.
New Zealand journal of geology and geophysics.
Lewis, K.B. 1979: Foraminifera on the continental shelf and slope off southern Hawkes
Bay, New Zealand. New Zealand department of scientific and industrial research
bulletin 163, 45 pp.

DNA studies of Recent foraminifera

Holzmann, M.; Pawlowski, J. 1997: Molecular, morphological and ecological
evidence for species recognition in Ammonia (Foraminifera). Journal of
foraminiferal research 27(4): 311-318.

Studies of former sea-levels using foraminifera

Cann, J.H., Belperio, A.P., Murray-Wallace, C.V. 2000: Late Quaternary
paleosealevels and paleoenvironments inferred from Foraminifera, northern
Spencer Gulf, South Australia. Journal of Foraminiferal Research 30: 29-53.
Hayward, B.W.; Grenfell, H.R.; Scott, D.B. 1999: Tidal range of marsh foraminifera
for determining former sea-level heights in New Zealand. New Zealand Journal
of Geology and Geophysics 42: 395-413.

Studies on biogeographic distribution of Recent foraminifera

Albani, A.D., 1970. A foraminiferal fauna from the Eastern Continental Shelf of
Australia. Contribution of the Cushman Foundation for Foraminiferal
Research, 21: 71-77.
Albani, A.D. and Yassini, I., 1989 - Taxonomy and Distribution of Shallow Water
Lagenid Foraminifera from the South Eastern Coast of Australia. Australian
Journal of Marine and Freshwater Research, 40 (4): 369-401.
Albani, A.D. and Yassini, I., 1994 - Taxonomy and distribution of the Family
Elphidiidae (Foraminiferida) from shallow Australian Waters. UNSW Centre for
Marine Science, Technical Contribution Series, No. 5, 54 pp.
Buzas, M.A.; Culver, S.J. 1989: Biogeographic and evolutionary patterns of continental
margin benthic foraminifera. Paleobiology 15(1): 11-19.
 33

Hayward, B.W.; Hollis, C.J.; Grenfell, H.R. 1997: Recent Elphidiidae

(Foraminiferida) of the South-west Pacific and fossil Elphidiidae of New
Zealand. Institute of Geological and Nuclear Sciences monograph 16, 166 pp.
Hayward, B. W.; Grenfell, H. R.; Reid, C. M.; Hayward, K. A. 1999: Recent New
Zealand shallow-water benthic foraminifera: taxonomy, ecologic distribution,
biogeography, and use in paleoenvironmental assessment. Institute of
Geological and Nuclear Sciences Monograph 21. 258 p.
Yassini, I. and Jones, B.G., 1995. Foraminiferida and ostracoda from estuarine and
shelf environments on the southeastern coast of Australia. Wollongong, NSW;
University of Wollongong: p.484

Recent planktonic foraminifera

Bé, A.W.H. 1977: An ecological, zoogeographic and taxonomic review of Recent
planktonic foraminifera. Pp. 1-100, In Ramsay, A.T.S. (ed..) “Oceanic
micropaleontology”. Academic Press, London.
Hayward, B.W. 1983: Planktic foraminifera (Protozoa) in New Zealand waters: a
taxonomic review. New Zealand journal of zoology 10: 63-74.

Application of fossil foraminifera to biostratigraphy

Albani, A.D., 1981. Sedimentary environments and Pleistocene chronology of the
Botany Basin, N.S.W., Australia. Geo-marine Letters, 1: 163-167.
Hornibrook, N.deB. 1982: Late Pliocene and Pleistocene Globorotalia
(Foraminiferida) from DSDP Leg 29, Site 284, Southwest Pacific. New Zealand
Journal of Geology and Geophysics 25: 83-99.
Hornibrook, N.deB.; Brazier, R.C.; Strong, C.P. 1989: Manual of New Zealand
Permian to Pleistocene foraminiferal biostratigraphy. New Zealand Geological
Survey paleontological bulletin 56, 175 pp.
Scott G.H.; Bishop S.; Burt B. 1990: Guide to some Neogene Globorotalids
(Foraminiferida) from New Zealand. New Zealand Geological Survey
paleontology bulletin 61. 135 pp.

Application of fossil foraminifera to basin analysis

Hayward, B.W. 1990: Use of foraminiferal data in analysis of Taranaki Basin, New
Zealand. Journal of foraminiferal research 20: 71-83.
 34

Hayward, B.W.; Brook, F.J. 1994: Foraminiferal paleoecology and initial subsidence
of the early Miocene Waitemata Basin, Waiheke Island, Auckland. New
Zealand journal of geology and geophysics 37: 11-24.
Hayward, B.W.; Buzas, M.A. 1979: Taxonomy and paleoecology of early Miocene
benthic foraminifera of northern New Zealand and the north Tasman Sea.
Smithsonian contributions to paleobiology 36: 1-154.
King, P.R.; Scott, G.H.; Robinson, P.H. 1993: Description, correlation and
depositional history of Miocene sediments outcropping along North Taranaki
Coast. Institute of Geological and Nuclear Sciences monograph 5, 199p.

Application of fossil foraminifera to sequence stratigraphy

Abbott, S.T. 1997: Foraminiferal paleobathymetry and mid-cycle architecture of mid-
Pleistocene depositional sequences, Wanganui Basin, New Zealand. Palaios 12:
267-281.
Haywick D.W.; Henderson R.A. 1991: A foraminiferal paleobathymetry of Plio-
Pleistocene cyclothemic sequences, Petane Group, New Zealand. Palaios 6:
586-599.
Kamp, P. J. J.; Journeaux, T. D.; Morgans, H. E. G. 1998: Cyclostratigraphy of
middle Pliocene mid shelf to upper slope strata, eastern Wanganui Basin (New
Zealand); correlations to the deep sea isotope record. Sedimentary Geology
117: 165-192.
Application of fossil foraminifera to plaoeclimate studies
Hornibrook, N.deB. 1992: New Zealand Cenozoic marine paleoclimates: A review
based on the distribution of some shallow water and terrestrial biota. Pp. 83-
106, In Tsuchi, R.; Ingle, J.C. (eds.) “Pacific Neogene environment, evolution
and events.” University of Tokyo Press.
Weaver, P. P. E., Neil, H.; Carter, L. 1997: Sea surface temperature estimates from
the Southwest Pacific based on planktonic foraminifera and oxygen isotopes.
Paleogeography, Paleoclimatology, Paleoecology 131: 241-256.

Application of fossil planktonic foraminifera to paleoceanography

Li, Q.; James, N.P.; McGowran, B.; Bone, Y.; Cann, J. 1998: Synergetic influence of
water masses and Kangaroo Island barrier on foraminiferal distribution, Lincoln
and Lacepede shelves, South Australia: A synthesis. Alcheringa 22: 153-176.
 35

Weaver, P.E.; Carter, L.; Neil, H.L. 1998: Response of surface water masses and
circulation to late Quaternary climate change east of New Zealand.
Paleoceanography 13: 70-83.

Application of fossil foraminifera to paleobiogeography and evolutionary history

Hayward, B.W. 1990: Taxonomy, paleogeography and evolutionary history of the
Bolivinellidae (Foraminiferida). New Zealand Geological Survey paleontological
bulletin 63, 132 pp.
Kennett, J.P. 1978: The development of planktonic biogeography in the Southern
Ocean during the Cenozoic. Marine Micropaleontology 3: 301-345.
Malmgren, B.A.; Kennett, J.P. 1981: Phyletic gradualism in a Late Cenozoic
planktonic foraminiferal lineage: DSDP Site 284: Southwest Pacific.
Paleobiology 7: 230-240.

Application of fossil foraminifera to studies of mass extinctions

Kaiho, K.; Arinobu, T.; Ishiwatari, R.; Morgans, H.E.G.; Okada, H.; Takeda, N.;
Tazaki, K.; Zhou, G.; Kajiwara, Y.; Matsumoto, R.; Hirai, A.; Niitsuma, N.;
Wada, H. 1996: Latest Paleocene benthic foraminiferal extinction and
environmental changes at Tawanui, New Zealand. Paleoceanography 11: 447-
465.
Strong, C.P. 2000. Cretaceous-Tertiary foraminiferal succession at Flaxbourne River,
Marlborough, New Zealand. New Zealand Journal of Geology and Geophysics.

Heavy liquid floating

Gregory M. and Johnston K.A. 1987: A nontoxic substitute for hazardous heavy
liquids - aqueous sodium polytungstate (3Na2WO4.9WO3.H2O) solution. New
Zealand Journal of Geology and Geophysics 30 (3): 317-320.
 36

APPENDIX - Geologic Time Scale

For easier reference a simplified geologic time scale is added.

Era Period Epoch m.y. (B.P.)

Recent
Quaternary 0.01
Pleistocene
1.8
Pliocene
Cainozoic 5
Miocene
23
Tertiary Oligocene
33
Eocene
55
Palaeocene
65
Cretaceous
140
Mesozoic Jurassic
190
Triassic
235
Permian
300
Carboniferous
345
Palaeozoic Devonian
390
Silurian
430
Ordovician
500
Cambrian 570

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