CHAPTER I

INTRODUCTION TO FORAMINIFERA

1.1 FORAMINIFERA

Herodotus, who observed the Nummulities in the rocks with which the Egyptian

pyramids were constructed, first recorded foraminifera in literature in the 5th century B.C.

However, it was until nearly 2,000 years later were they recognized as being the fossil

remains of organisms. This was by Agricola in 1558 A.D. The smaller foraminifera were

first described by Beccarius in 1731. Dujardin first demonstrated the protozoan nature of

these organisms in 1835.

The first larger scale systematic work was that of d’Orbigny (1826), in which 5

families, 52 genera and 544 species were recognized. Since he defined genera and species

within narrow limit and cited their geological occurrence as well, this early taxonomic

work also represents the earliest biostratigraphic application of this group of organisms.

While the early continental workers concentrated on stratigraphic application of

the foraminifera and faunal description (e.g. d’Orbigny, Reuss, Terquem, Berthelin), the

English work, in general, concentrated on the morphological studies and description of

recent faunas (e.g., Brady, Carter, Williamson, Heron-Allen, Earland).

Foraminifera are micro-unicellular organisms. They are almost entirely marine

animals though a few live in brackish and even fresh water. These organisms build a shell

and for paleontologists, the characteristics of the shell are the primary feature which can

be used to distinguish one species from another and hence, to use these distinction to

form interpretation of time or environment.

Foraminifers constitute the most diverse group of shelled microorganisms in

modern oceans. Shells (usually called “tests”) of extant foraminifers have been noticed in

shore sands since the 17th century. An unusual early find was by the great microscopist,
Antonie van Leeuwenhoek who, in 1700, wrote about foraminifer shells “no bigger than

a coarse sand grain” in the stomachs of shrimps, and described the specimens as “no

bigger than a coarse sand grain” in stomachs of shrimps, and described the specimens as

“very little small shells”.

The biology of foraminifers has not been examined as intensively as that of some

protoctistan cousins (e.g. ciliates); apparently, the shell has been somewhat of a problem

in cellular studies. To most students and researchers, however, the most significant aspect

of foraminiferal biology is the production of an amazing variety of shells, matched in no

other class of organisms.

On the basis of the morphology of foraminiferal tests, 15 extant foraminiferal

orders can be recognized. In 7 orders, the test is made up of secreted calcite; in others, the

species secrete aragonite or opaline silica, or construct their tests with organic matter of

foreign particles.

The ecology of foraminifera became a major area of study in the second half of

the 20th century, and the first text on this subject was by Fred B. Phleger (1960). In the

past 30 years, research in this field has increased tremendously, mainly because of the

realization that modern foraminiferal distributions are likely to provide reliable clues for

the understanding of marine environmental changes in the geological and, in some cases,

the historical past.

The advantage of using foraminifers in paleoecology lies in their significant

numerical density in diverse marine sediments, and in the excellent preservation of their

tests. Both of these factors are partly related to their small size. Some species are,

however, better preserved than others, causing a taphonomic bias even in relatively recent

sedimentary records.
 At the sediment-water interface, many foraminifers are neither obligate

epibenthos (living on the sediment) nor obligate endobenthos (living within the

sediment), because they can move up and down the sediment column in search of food,

and thus have variable microhabitats (Linke and Lutze, 1993). Some truly epibenthic

species (free or attached) live on substrates above the sediment surface, whereas some

truly endobenthic species live several centimeters down in the sediment. Such deep

endobenthic species may have to cope up with a severe shortage of oxygen in the

sediment pore water. But recent research has shown that many foraminiferal species can

live in extremely oxygen-poor water in varying depths of the sea.

Due to the paucity of observations on the preferred depths of living individuals

within the substrate, microhabitats of most sediment-dwelling species are unknown. In

many coastal areas, especially in industrialized countries, pollution has severely affected

foraminiferal microhabitats. Studies on the effects of various forms of pollution on

foraminifers have shown that many species can serve as indicators in pollution

monitoring programs.

Many foraminiferal species have adapted to extreme natural environments, such

as habitats of high salinity, areas near hydrothermal vents (Jonasson et al., 1995),

bacterial mats at hydrocarbon vents (Sen Gupta et al., 1997), or in silled basins (e.g.

Bernhard et al., 1997), and pack ice (planktonic species).

Modern planktonic foraminifera, because of their enormous population sizes,

produce a significant amount of oceanic carbonate. There is a much larger proportion of

taxa with algal symbionts within this group than within the benthic group. Environmental

controls on planktonic species are much better understood than those on benthic species,

because the only major variables are temperature, salinity, and productivity of the surface

layer of sea water, mainly in the upper 100 m.

 Planktonic species are distributed in a few, large latitudinal provinces, with some

showing bipolar distributions; many species can be used as tracers of ocean currents

(Oberhänsli, 1992). The present-day temperature limits of the biogeographic provinces

have been of great use in estimating Quaternary se-surface paleo-temperatures from the

fossil record of extant species.

The extraordinary progress of foraminiferal test geochemistry in recent years has

been in the context of major questions of paleo-oceanography that relate to the nature and

movement of past water masses, at the sea surface and in the water column. The

principles behind these inquiries have to be established by analyses of modern

foraminiferal tests – both planktonic and benthic. Culturing studies have become

especially significant in this research, because of the necessity of understanding species-

specific “vital effect” in the incorporation of trace elements and the acquiring of oxygen

and carbon isotope signatures in the test.

1.2 LIFE HISTORY AND ECOLOGY OF FORAMINIFERA

Most of the estimated 4,000 living species of foraminifera live in the world's

oceans. Of these, 40 species are planktonic, that is they float in the water. The remaining

species live on the bottom of the ocean, on shells, rock and seaweeds or in the sand and

mud of the bottom. In places, foraminifera are so abundant that the sediment on the

bottom is mostly made up of their shells. For example, the pink sands of Bermuda get

their color from the shells of a foraminiferan called Homotrema rubrum which has pink

to red-colored shells. Far from land in the deep sea, where little material comes from

erosion of the land, the bottom sediment is made up mainly of shells of planktonic

organisms, especially foraminifera.

The degree to which deep sea sediments in an area are composed of one or more

of biogenic types depends on the organic productivity of various organisms in the surface
water, the degree to which the skeletal remains are re-dissolved by sea water while

settling to the bottom, and the rate of sedimentation of other types of sediment material.

Where the sediments are composed largely of a single type of biogenic material, it is

often referred to as an ooze, after its consistency in the place on the ocean floor. Thus,

depending on the organism, it may be called a foraminiferal ooze or Globigerina ooze,

after one of the most abundant planktonic genera, Globigerina.

Foraminifera are found in all marine environments, from the intertidal to the

deepest ocean trenches, and from the tropics to the poles, but species of foraminifera can

be very particular about the environment where they live. Some are abundant only in the

deep ocean; others are found only in brackish estuaries or salt marshes along the shore,

and most live at certain depths and water temperatures in between.

Foraminifera are an important part of the marine food chain. On the continental

shelf there can be tens of thousands of living individuals per square meter of the ocean

bottom. Many larger animals (including snails, sand dollars, and fish) eat foraminifera,

and some are very selective about which species they eat.

As different species of foraminifera are found in different environments,

paleontologists can use their fossils to determine past environments. If a sample of fossil

foraminifera contains many living species, the present-day distribution of those species

can be used to infer the environment there when the fossils were alive. Even when

samples contain all or mostly extinct species, such data as species diversity, the relative

numbers of planktonic and benthic species (planktonic : benthic ratio), and the ratios of

different shell types are used to infer past environments.

In addition to using species distributions (whether directly or through diversity

and other ratios) to study past environments, the chemistry of the shell can tell us about

the chemistry of the water in which it grew. Most importantly, the ratio of stable oxygen
isotopes depends on the water temperature, because warmer water tends to evaporate off

more of the lighter isotopes. Studies of stable oxygen isotopes in planktonic and benthic

foram shells from hundreds of deep-sea cores worldwide have been used to map past

water temperatures. These data help in understanding how the climate has changed in the

past, and thus how it may change in the future.

1.3 BRIEF HISTORY OF FORAMINIFERAL STUDIES

It is interesting to note that foraminifera received their name before their nature

was understood (Lister, 1903). In the latter half of the 18th century, most foraminifers

were described as cephalopods, particularly as Nautilus. The original descriptions of

about 140 such false nautiluses named between 1758 and 1819 are listed in the Catalogue

of Foraminifera (Ellis and Messina, 1940 and later). Many of the species and varieties

were, however, well illustrated by their authors, and it is possible to place them under

currently accepted generic labels.

The best known phase of foraminiferal taxonomy in the 18th century is the

Testacea Microscopica of Fichtel and Moll (1798), in which 47 extant and fossil taxa

were assigned linnean names under the single generic heading of Nautilus; 18 separate

genera, all extant, are now recognized within this group (Rögl and Hansen, 1984).

One of the greatest works on modern foraminifera was H. B. Brady’s magnificent

monograph (1884) on the material collected during the Challenger Expedition of 1873-

1876. Brady’s illustrations are still used extensively for species identifications; names

may have changed because of questions of priority, but by and large, the internal

consistency of nomenclature is impressive (Barker, 1960; Jones, 1994). Brady’s views on

foraminiferal systematics, forcefully expressed in the Challenger monograph, strongly

influenced researchers in the next five decades.

1.4 CLASSIFICATION OF FORAMINIFERA

As a result of 500 ma of evolution in numerous independent lineages, there now

exists a vast array of foraminiferal morpho-types. Loeblich and Tappan (1964) estimated

that there are close to 100 families, over 1,200 genera, and some 27,000 species of

foraminifera described in the literature on foraminifera. To organized and categorize this

vast variety of forms, over 35 schemes of classification have been proposed, some dating

back to the very first publication on foraminifera. These classifications are summarized in

Loeblich and Tappan (1964).

Early classifications were necessarily based on gross feature of test morphology

as well as on limited faunas. More recent classifications attempt a ‘natural’ classification

based on as many criteria as possible, usually in following hierarchical order:

1. Wall composition and microstructure

2. Chamber arrangement and septal addition

3. Apertural character and modification

4. Chamber forms

5. Life habits and habitats

6. Protoplasm characteristics

7. Ontogenic changes

8. Reproductive processes

9. Evolutionary range

The identification and separation of foraminiferal species, genera, and higher taxa

solely by aspects of test morphology, especially chamber arrangement, did not change

with the recognition by Felix Dujardin that the foraminiferal body was simply a jelly-like

mass (“sarcode”) with pseudopodia. The classification in vogue at this time (d’Orbigny,

1826) underwent only minor modifications after d’Orbigny’s acceptance of Dujardin’s

conclusion that foraminifers were unicellular organisms, and not cephalopods. By 1852,

d’Orbigny had recognized a total of 72 foraminiferal genera.

Although wall characters were considered a significant criterion in generic

taxonomy, d’Orbigny (1852) placed both calcareous and agglutinated foraminiferal

genera in the same family, because of some similarity in chamber arrangement. For

example, the agglutinated form of Clavulina and the calcareous forms Bulimina and

Globigerina were placed under the family Tubinoidaea. This practice of placing

agglutinated and calcareous genera (especially those with serial chamber arrangements)

within one family continued well into the 20th century.

Cushman’s first classification (1925) was simply an adaptation of Brady’s 10

families with some changes in the naming of subfamilies. The big expansion came with

his 1927 and 1928 classifications, in which he increased the number of families to 45,

without proposing superfamilies or orders. Apart from test morphology, Cushman’s new

classification was based upon the known geological history of the genera, upon the

phylogenetic characters through a study of very much fossil material from all the

continents, and finally by a study of the ontogeny in very many microspheric specimens,

which show the relationships much more definitely than do megalospheric specimens of

the same species (Cushman, 1933).

Within about 20 years after its publication, Cushman’s 1927 classification became

the standard classification, adopted by workers on the foraminifera throughout the world.

A contributing factor was Cushman’s reputation as the foremost specialist in the use of

fossil foraminifers in petroleum exploration. Biological research on foraminifers,

however, had not progressed much beyond what it was at the beginning of the 20th

century.
 Several classifications developed in the two decades following Cushman’s last

major taxonomic work (1948) introduced a suprafamilial hierarchy within the order

Foraminifera, but none received a wide acceptance. This scene changed with the

appearance of Loeblich and Tappan’s first book on foraminiferal systematics (1964).

Meanwhile, a landmark paper on foraminiferal wall structure (Wood, 1949) focused

attention on characters of calcitic foraminiferal walls that had been more or less ignored

until this time.

Following Galloway (1933) and Cushman (1945), Loeblich and Tappan (1964)

argued that the same chamber arrangement and form of test may have developed in

independent lineages by parallel evolution, without indicating inter-relationship of the

similarly shaped tests. Thus, in the descending hierarchy of classification, the starting

point was the nature of the test wall. The wall composition and texture formed the basis

for the separation of the five suborders: 1. Organic wall for the Allogromiina, 2.

Agglutinated wall for the Textulariina, 3. Calcareous, microgranular wall for the extinct

Fusulinina, 4. Calcareous, porcelaneous wall for the Miliolina, and 5. Calcareous, hyaline

wall for the Rotaliina.

Several modifications to their 1964 classification were proposed by Loeblich and

Tappan in later years (Loeblich and Tappan, 1974, 1984; Tappan and Loeblich, 1982),

culminating a large compendium (Loeblich and Tappan, 1987; not 1988), as in some

citations in the literature, with descriptions of 2,446 genera (Haman, 1988), including the

878 extant genera (Tappan and Loeblich, 1988). A brief, but significant, revision of the

taxonomic hierarchy (orders to sub-families) was published in 1992. The 1987

classification included 12 suborders. In the 1992 revision, 10 of these were changed to

orders, 4 new orders were recognized, and the taxon Foraminifera (labeled

“Foraminiferea” after Lee, 1990) was raised to the rank of class.

 With 1,400 genera, and 30,000 species (including 4,500 still in existence), the

Order Foraminiferida accounts for half the known protozoa. Their classification is

difficult and is based on the characteristics of the test. The most utilized classification has

been that by Loeblich and Tappan (1987).

1.5 EVOLUTION OF FORAMINIFERA THROUGH TIME

The foraminifera evolved in the Cambrian (e.g. Bathysiphon), as forms with

unilocular agglutinated tests. Microgranular, calcareous tests emerged in the Silurian and

pseudofibrous types in the Devonian. An important event took place at the Devonian-

Carboniferous boundary; this was the development of partition yielding multilocular tests

with periodic growth. Not long afterwards, during the Carboniferous, porcelaneous tests

appeared together with morphological dimorphism and trochospiral coiling; these

occurred in the agglutinated Trochammina and thelate calcareous Tetrataxis. The

Endothyridae and the Fusulinidae proliferated in the Paleozoic before disappearing on the

threshold of the Mesozoic.

Although poor in foraminifera, the Triassic saw considerable renaissance which,

during the Jurassic, took the form of the development of the true Miliolodae, the first

hyaline tests to be fibroradial, perforate and lamellar (Nodosariidae), and planktonic

forms (Globigerina).

During the early Cretaceous, some species began to population lagoons.

Diversification continued in the Cretaceous with the acquisition of a canal system

(Rotaliidae), the proliferation of planktonic forms (Globotruncana) and the larger benthic

group (Orbitolinidae, Alveolinidae and Orbitoididae).

The beginning of the Cenozoic withessed the sudden extinction of many genera,

both benthic (Orbitoides) and planktonic (Globotruncana). This was soon followed by a
new and important phase of diversification affecting old group (Alveolinidae,

Orbitoididae, and planktonic forms) and engendering new ones (Nummulitidae).