http://oceanworld.tamu.edu/students/forams/forams1.htm

Introduction

Foraminifera, or informally "forams", are tiny marine organisms. They are fun to study because they are not only an important part of the marine food web but also because they provide big clues to scientists trying to understand our earth, ocean and atmosphere.

This series of web pages explores various aspects of forams. In What is a Foram? you will find out what living forams are like - what they look like, what they eat and where they live. In Foram Test Construction we will discuss how foram tests (shells) have changed through time and share some cool images of the four test types.

Foram Evolution focuses on how forams have evolved over the last 500 million years and Why are Forams Important? outlines the unique characteristics of forams that make them so useful to scientists. Our Forams for Correlation and Forams Highlight Pollution subtopics describe ways that forams are routinely being used by scientists today; the first looks at how forams are used in the Integrated Ocean Drilling Program and in the Petroleum Industry, while the second steps through how forams are aiding in pollution studies of Biscayne Bay, Florida and in Long Island Sound. To see the full organization of the Forams Unit, select Foram Topic Map.

If you are interested in research and would prefer to access raw samples, data or images go to Foram Data Sets.

Rosalina globularis in darkfield illumination. (Photo courtesy the Foram Gallery by Wim van Egmond and Brian Darnton)

Take a quick look at the photo above. This is a foram test without the organism. Think of it like a shell you pick up along the beach, except this test is very small, about the size of a grain of sand. What are those small bumps? What might the whole organism have looked like? Did they all look the same? What did they eat and where did they live?

WHAT is a FORAM ?

Foraminifera are single-celled organisms that have inhabited all types of marine environments for millions of years.  Their survival is due in part to their ability to produce a protective outer shell, called a test.  A test surrounds and protects them, and is composed of material that ranges from calcium carbonate to sand grains.  Forams are categorized by their size and whether they are planktonic or benthic.  The abundance, wide distribution, and sensitivity to their environment make these creatures unique and extremely helpful in studies of the oceans.

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Globigerinoides ruber - fossil shell of a planktonic foraminifera.(Photo courtesy Christa Farmer of Columbia University)

Benthic? Planktonic? Gesundheit!BENTHIC VS. PLANKTONIC

Benthic foraminifera are commonly refered to as the bottom dwellers.  They live all along and beneath the ocean floor in the sediments.  Benthic organisms live in a wide array of environments, ranging from marshes to abyssal plains.   They are able to move and feed by use of pseudopodia. The type of pseudeopodia varies for each species.  They are excellent indicators of ocean depth and serve as the primary biostratigraphic indicators for paleontologists.  In just a handful of sediment, thousands of forams can be found.  Their small size is key in how important they are to research.  Planktic foraminifera live in the upper zone of the ocean.  These creatures are distributed worldwide, but found only in the open ocean.  When they die, they settle to the bottom of the ocean.  Planktonic forams are indicators of ocean currents and climates. The planktic and benthic forams can easily be seperated, because specific forams only live in special conditions and environments.

Bolivina subaenariensis - foraminifera approximately 1mm long.(Photomicrograph courtesy B. Sen Gupta, Louisiana State University)

How do forams evolve?FORAM EVOLUTION

Hantkenina alabamensis - planktonic foraminifera. (Photo courtesy J. H. Lipps of University of California, Davis)

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Forams are similar to most species in the respect to how they evolved.  They would experience explosive radiation and evolution, followed by mass extinction.  The first known species of forams are from the Cambrian.  This benthic foram was simple agglutinated tubes with only a single chamber.  During the Devonian , the forams evolved to have multiple chambers and a few were calcareous.  Calcareous forams began to really radiate in the Carboniferous.  All Paleozoic forams were benthic.  Planktonic forams did not evolve until the Mesozoic, and radiated several times during the Cenozoic.  Like most organisms of the time, the mass extinction during the Permian killed thousands of species of forams.

Pangea - This is how the Earth looked during the mass extinction of the Permian. (Image courtesy Dr. Christopher Scotese)

CORRELATIONS

The mass extinctions of forams correlates with other major events in the Earth's history.  The Earth has a cyclic pattern that goes from Ice Ages to normal.  In times that are not associated with ice ages, the polar caps are not completely frozen and this increases the worldwide water levels.  This change in water level, causes the depth of the ocean to change also.  The fossil record also indicates how the foram's environment changes throughout these times.  Forams must migrate or become extinct if it lives during a time when it's environment changes.  Glaciers melting during the Permian is a probable cause for the extinction of forams, because of the rapid warming that was associated with this time.

Benthic forams secrete their shells from calcium, magnesium, and oxygen dissolved in sea water. (Photo courtesy USGS)

WHAT CAN THEY TELL US ?

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Orbulina universa - sand-sized central sphere foraminifera surrounded by delicate spines. (Photo courtesy Dr. Howard Spero of UC Davis)

Several different groups are interested in fossils for different reasons. Petroleum geologists often work with paleontologists, because a paleontologist can determine by the type of forams that are in a rock layer whether there will be oil present. Forams are also used by geochemists. These scientists are able to determine the fossil's age by ratio-isotope studies. This is done by measuring the amount of element, such as carbon, that remains in the fossil. Whether it is students in a classroom or a professor in a lab, the small size and abundance of forams make them useful tools for anyone to work with.

What is a Foram?Forams are single-celled organisms that typically live in the ocean and produce a mineralized test (shell). Foram tests are generally composed of secreted calcium carbonate (CaCO3), but less commonly they may be composed of organic material or cemented particles scavenged from the sea floor.

Forams produce tests in a range of shapes and sizes. Some tests are simple, single-chambered forms like the one shown in the photo below, while others are multi-chambered and elaborately structured. Typically they are microscopic in size, and generally range from 0.1 to 1 mm. (approximately the size of a grain of sand or smaller). However, in the geologic past, forams with test diameters greater than 10 cm. (4 in.) were not unusual.

The test houses and protects the organism. Thin organic filaments called rhizopodia (shown as white radiating threads in the photo above) extend through perforations in the test. Do you remember seeing bumps on the test pictured on the Foram Introduction page? Those are pores through which rhizopodia once extended. These rhizopodia help the foram eat, build its test, and depending on the type of foram, rhizopodia help it to move or to stay attached to the sea floor.

Forams eat a variety of foods including organic material dissolved in sea water, algae, bacteria and the larvae of small animals such as crustaceans or fish. Some foram species even host symbiotic algae, which provide the forams with steady nourishment in waters poor in other food sources. By the way, both species pictured on this page host symbiotic algae. Algae-hosting species are most common in latitudes near the equator where other food sources are often in short supply. In their turn, forams are eaten by fish and other small marine animals, and thus form an important part of the marine food web.

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Globigerinoides sacculfier, a living multi-chambered planktonic foram. (Photo used by permission, Dr. David A. Caron, University of Southern California)

Forams are distributed throughout the world's oceans from the North Pole to the South Pole. They are found at all depths and can tolerate the whole range of salinity, temperature, and light conditions. Almost all varieties prefer the marine environment, although a few unusual species are found in fresh water.

There are two major classes of forams,

1. benthic - those that live on top of or just within the sea floor, and 2. plankontic - those that float freely in shallow, sunlit water. The vast majority of

present-day species are benthic.

Foram Test ConstructionAs we mentioned on the previous web page What is a Foram? forams are single-celled organisms with a protective test (shell). What is the test made from and how does the foram make it? There are two basic kinds of tests, soft tests made from an organic material called tectin (a complex carbohydrate plus protein material) and hard tests made from minerals. Hard, mineralized tests are much more common.

The evolution of a hard outer shell likely provided forams with additional protection against predators, physical damage, chemical changes, and as a means to control buoyancy. Mineralized foram tests are sorted into four major classes based on the method the foram used to construct them.

1. Agglutinated 2. Microgranular 3. Porcellaneous 4. Hyaline

1. Agglutinated - This test style was the first to evolve (more than 500 million years ago) and consists of tiny cemented grains gathered from the sea floor. Particles are glued to a tectin base with an organic, calcite or iron-bearing cement. Remember the organic tests we first talked about? This is simply an organic test with particles glued on top. At times these particles appear to have been gathered at random and at other times the grains are similar in size, composition or shape.document.doc 5 of 38

More images of agglutinated forams

Bigenerina nodosaria, Agglutinated test from Puerto Rico, (SEM) image -120X magnification

(Credit: Heidi Crevison Souder; used by permission from Foraminifers as Bioindicators website, University of South Florida, St. Petersburg)

2. Microgranular - Test walls are composed of tiny, closely-packed grains of calcium carbonate, but with no obvious cement. For many years this group was lumped into the agglutinated class because it was thought that these tests were built from scavenged grains. In fact, some species do have some agglutinated grains thereby adding to the confusion. However, scientists are now convinced this group of forams actually secrets the tiny grains.

Image of microgranular forams

3. Porcellaneous - This test is composed of microscopic calcite needles formed inside the foram and then moved to the outside of the cell. Tests are generally white in color, but may be pink or colorless with an appearance similar to porcelain, hence the name "porcellaneous". Test walls are generally built of three layers - a central layer sandwiched between two outer layers. Calcite needles in the middle layer are arranged in a poorly-organized mesh-like network, while needles in the outer layers are arranged parallel to one another, providing the outside of the test with a smooth surface.

Images of porcellaneous forams

Discorbis mira, Hyaline test from the Florida Keys, (SEM) image -150X magnification

(Credit: Heidi Crevison Souder; used by permission from Foraminifers as Bioindicators website, University of South Florida, St. Petersburg)

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4. Hyaline - To build this type of test, the foram secrets one or more layers of calcium carbonate over soft tectin material that serves as a template for the final shape. Rhizopodia, thin organic filaments connected to the foram, assist with hyaline test construction by delivering liquid calcium carbonate to different patches of the exposed tectin surface. The patches of calcium carbonate eventually merge and harden into a solid shell. Hyaline tests are often translucent to light, and commonly have a glassy or milky iridescent appearance.

More images of hyaline forams

Did you know that forams first appeared on Earth more than 500 million years ago?

Foram EvolutionForams live today in all the world's oceans, but they first appeared in the rock record approximately 525 million years ago. Between that long ago time and today, the size, shape and construction of foram tests changed significantly. Tests might be single-chambered or multi-chambered, right coiling or left coiling, larger or smaller, or they may be constructed from scavenged grains or self-produced CaCO3, depending on the conditions and time period in which the foram lived.

Hantkenina alabamensis - planktonic foraminifera. (Photo credit: J. H. Lipps of University of California, Davis, used by permission)

Traditionally, forams have been grouped into species on the basis of their characteristic test shape; and their distribution has been studied. Scientists discovered that some species existed only during narrow geologic time bands, while other species survived much longer. Some were restricted in their geographic distribution, while others were global in extent; and some species were found exclusively in shallow water environments, while others preferred deeper water.

Over time, scientists have painstakingly pieced together the geologic time period, geographic distribution and preferred environment for many individual foram species, and from this work some major events in foram evolution are evident.

Major Events in Foram EvolutionEarly Cambrian (~525 million years ago)

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Forams first appeared in the rock record. Earliest specimens were agglutinated, single-chambered varieties; meaning that the forams created their shell by cementing together tiny particles gathered from the sea floor. All species were benthic (bottom-dwelling); however some benthic species moved about on the sea floor while others were stationary either by attaching themselves to the bottom or by burrowing into it.

Late Cambrian (>500 million years ago)

Multi-chambered varieties first evolved.

Devonian (>360 million years ago)

Microgranular and porcellaneous (biomineralized) calcareous tests first evolved.

Middle Pennsylvanian (~308 million years ago)

Forams evolved a hyaline (glassy) calcareous test wall. Additionally, larger foram species first appear, possibly to serve as hosts to symbiotic algae.

End Permian (~250 million years ago)

Mass extinction of most foram species including the large Fusilinids. This extinction is believed to be the largest in Earth's history with 90-95% of all marine species becoming extinct.

Early Jurassic (~183 million years ago)

Planktonic (floating) forams first appear in the rock record. Until this time, all species had been benthic.

Middle Cretaceous (~112 million years ago)

Geographic distribution of planktonic forams begins to expand rapidly.

End Cretaceous (~65 million years ago)

Decrease in planktonic diversity, and extinction of many planktonic foram species. Survivors were generally smaller in size than extinct species. Benthic species were not substantially impacted.

End Paleocene (~55 million years ago)

Extinction of approximately one-half of deep water benthic foram species with an associated decrease in deep water benthic diversity of approximately 30-50%.

Late Eocene to Early Oligocene (~30-39 million years ago)

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Numerous smaller extinctions and appearances of benthic foram species throughout this time period, with an marked extinction of many planktonic foram species around 37 million years ago.

Middle Miocene (~12-19 million years ago)

Major planktonic and benthic foram species composition and abundance changes are documented for the middle Miocene. Modern benthic foram varieties evolve.

Today

More than 10,000 foram species are living today. The vast majority of these are benthic; only about 40-50 species are planktonic. Benthic forams today are nearly identical to the suite that evolved in the middle Miocene

Why are Forams Important?

Foram tests have six key characteristics that make them useful to scientists.

1. Small

Foram tests are typically about the size of a grain of sand or smaller. Scientists who study marine geology often only have small samples of material to work with, such as from a piston core. Consequently, in these small samples a small fossil is much more likely to be present and undamaged than a big fossil would be.

2. Abundant and geographically distributed

Foram tests are common constituents of marine sediments, and they have been common for a long time. In some instances, tests can accumulate in such great numbers (sometimes in thicknesses of up to 1km!) that the marine rock is named after them. These rocks are called foraminiferal oozes. Even if they are not found in this abundance, a few tests in a single sample can help guide scientists in their work.

Cluster of planktonic forams (Credit: NOAA Paleoclimatology Program, US Department of Commerce)

3. Test shape and size differs through time

Foram tests have differed through the ages. Scientists built on this fact, and learned to tie specific test shapes to specific time periods. This knowledge permits them to estimate a sample's geologic age without having to perform expensive and time-consuming absolute age-dating analyses. In addition, scientists have also tied specific test shapes to water depth.

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Some tests are found only in shallow water environments; others in moderate depths, and still others are found only in deep water environments.

4. Existed for more than 500 million years

Forams first appeared in the rock record during the Cambrian geologic time period and they are thriving today. Because their record is so long in duration, it is easier to recognize changes in test shape, species abundance and species distribution.

5. Short reproductive cycles

The reproductive cycle for modern forams is relatively short (6 months to 1 year), making them particularly useful to scientists looking for pollution-related growth deformities.

6. Trace element chemistry preserved in test

As forams construct their test, trace elements may be scavenged from the water column and incorporated into the test wall. These elements tell scientists about ocean water chemistry and temperature at the time the test was formed.

How do marine geologists use forams to determine geologic age?

Go to Forams for Correlation to find out!

ForaminiferaForaminifera

Fossil range: 600 Ma PreЄ

↓

Precambrian - Recent

Live Ammonia tepida (Rotaliida)

Scientific classificationDomain: EukaryotaKingdom: RhizariaSuperphylum: RetariaPhylum: Foraminifera

d'Orbigny, 1826

Orders

AllogromiidaCarterinidaFusulinida - extinctGlobigerinidadocument.doc 10 of 38

Involutinida - extinctLagenidaMiliolidaSilicoloculinidaSpirillinidaTextulariidaincertae sedis   Xenophyophorea   Reticulomyxa

The Foraminifera, ("Hole Bearers") or forams for short, are a large group of amoeboid protists with reticulating pseudopods, fine strands of cytoplasm that branch and merge to form a dynamic net. They typically produce a test, or shell, which can have either one or multiple chambers, some becoming quite elaborate in structure. These shells are made of calcium carbonate (CaCO3) or agglutinated sediment particles. About 275,000 species are recognized, both living and fossil. They are usually less than 1 mm in size, but some are much larger, and the largest recorded specimen reached 19 cm.

Although as yet unsupported by morphological correlates, molecular data strongly suggest that Foraminifera are closely related to the Cercozoa and Radiolaria, both of which also include amoeboids with complex shells; these three groups make up the Rhizaria. However, the exact relationships of the forams to the other groups and to one another are still not entirely clear.

Living foramsModern forams are primarily marine, although they can survive in brackish conditions. A few species survive in fresh water and one even lives in damp rainforest soil. They are very common in the meiobenthos, and about 40 morphospecies are planktonic. This count may however represent only a fraction of actual diversity, since many genetically discrepant species may be morphologically indistinguishable. The cell is divided into granular endoplasm and transparent ectoplasm. The pseudopodial net may emerge through a single opening or many perforations in the test, and characteristically has small granules streaming in both directions.

The pseudopods are used for locomotion, anchoring, and in capturing food, which consists of small organisms such as diatoms or bacteria. A number of forms have unicellular algae as endosymbionts, from diverse lineages such as the green algae, red algae, golden algae, diatoms, and dinoflagellates.

Some forams are kleptoplastic, retaining chloroplasts from ingested algae to conduct photosynthesis.

The foraminiferal life-cycle involves an alternation between haploid and diploid generations, although they are mostly similar in form. The haploid or gamont initially has a single nucleus, and divides to produce numerous gametes, which typically have two flagella. The diploid or schizont is multinucleate, and after meiosis fragments to produce new gamonts. Multiple rounds of asexual reproduction between sexual generations is not uncommon in benthic forms. Foramanifera typically live for about a month.

Tests

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Foraminiferan tests (ventral view)

Fossil nummulitid forams showing microspheric and megalospheric individuals; Eocene of the United Arab Emirates; scale in mm.

The form and composition of the test is the primary means by which forams are identified and classified. Most have calcareous tests, composed of calcium carbonate.[4] In other forams the test may be composed of organic material, made from small pieces of sediment cemented together (agglutinated), and in one genus of silica. Openings in the test, including those that allow cytoplasm to flow between chambers, are called apertures.

Tests are known as fossils as far back as the Cambrian period,[8] and many marine sediments are composed primarily of them. For instance, the limestone that makes up the pyramids of Egypt is composed almost entirely of nummulitic benthic foraminifera.[9] Production estimates indicate that reef foraminifera annually generate approximately 43 million tons of calcium carbonate and thus play an essential role in the production of reef carbonates.[10]

Evolutionary significanceDying planktonic foraminifera continuously rain down on the sea floor in vast numbers, their mineralized tests preserved as fossils in the accumulating sediment. Beginning in the 1960s, and largely under the auspices of the Deep Sea Drilling, Ocean Drilling, and International Ocean Drilling Programmes, as well as for the purposes of oil exploration, advanced deep-sea drilling techniques have been bringing up sediment cores bearing foraminifera fossils by the millions. The effectively unlimited supply of these fossil tests and the relatively high-precision age-control models available for cores has produced an exceptionally high-quality planktonic foraminifera fossil record dating back to the mid-Jurassic, and presents an unparalleled record for scientists testing and documenting the evolutionary process. The exceptional quality of the fossil record has allowed an impressively detailed picture of species inter-relationships to be developed on the basis of fossils, in many cases subsequently validated independently through molecular genetic studies on extant specimens.

Uses of foramsBecause of their diversity, abundance, and complex morphology, fossil foraminiferal assemblages are useful for biostratigraphy, and can accurately give relative dates to rocks. The oil industry relies heavily on microfossils such as forams to find potential oil deposits.[12]

Calcareous fossil foraminifera are formed from elements found in the ancient seas they lived in. Thus they are very useful in paleoclimatology and paleoceanography. They can be used to reconstruct past

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climate by examining the stable isotope ratios of oxygen, and the history of the carbon cycle and oceanic productivity by examining the stable isotope ratios of carbon;[13] see δ18O and δ13C. Geographic patterns seen in the fossil records of planktonic forams are also used to reconstruct ancient ocean currents.[citation needed] Because certain types of foraminifera are found only in certain environments, they can be used to figure out the kind of environment under which ancient marine sediments were deposited.[citation needed]

For the same reasons they make useful biostratigraphic markers, living foraminiferal assemblages have been used as bioindicators in coastal environments, including indicators of coral reef health.[citation needed] Because calcium carbonate is susceptible to dissolution in acidic conditions, foraminifera may be particularly affected by changing climate and ocean acidification.[citation needed]

Foraminifera can also be utilised in archaeology in the provenancing of some stone raw material types. Some stone types, such as chert, are commonly found to contain fossilised foraminifera. The types and concentrations of these fossils within a sample of stone can be used to match that sample to a source known to contain the same 'fossil signature'.

http://www.ucmp.berkeley.edu/fosrec/Wetmore.html

WHAT ARE FORAMINIFERA?Foraminifera (forams for short) are single-celled organisms (protists) with shells or tests (a technical term for internal shells). They are abundant as fossils for the last 540 million years. The shells are commonly divided into chambers that are added during growth, though the simplest forms are open tubes or hollow spheres. Depending on the species, the shell may be made of organic compounds, sand grains or other particles cemented together, or crystalline CaCO3 (calcite or aragonite).

Fully grown individuals range in size from about 100 micrometers to almost 20 centimeters long. Some have a symbiotic relationship with algae, which they "farm" inside their shells. Other species eat foods ranging from dissolved organic molecules, bacteria, diatoms and other single-celled algae, to small animals such as copepods. They catch their food with a network of thin pseudopodia (called reticulopodia) that extend from one or more apertures in the shell. Benthic (bottom-dwelling) foraminifera also use their pseudopodia for locomotion.

WHERE DO THEY LIVE?There are an estimated 4,000 species living in the world's oceans today. Of these, 40 species are planktonic, that is they float in the water. The remainder live on or in the sand, mud, rocks and plants at the bottom of the ocean. 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 environmentin which they live. Some are abundant only in the deep ocean, others are found only on coral reefs, and still other species live only in brackish estuaries or intertidal salt marshes.

Foraminifera are among the most abundant shelled organisms in many marine environments. A cubic centimeter of sediment may hold hundreds of living individuals, and many more dead shells. In some environments their shells are an important component of the sediment. For example, the pink sands of some Bermuda beaches get much of their color from the pink to red-colored shells of a foraminiferan. In regions of the deep ocean far from land the bottom is often made up almost entirely of the shells of planktonic species.

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WHY ARE THEY IMPORTANT?The study of fossil foraminifera has many applications beyond expanding our knowledge of the diversity of life. Fossil foraminifera are useful in biostratigraphy, paleoecology, paleobiogeography, and oil exploration.

BIOSTRATIGRAPHY

Foraminifera provide evidence of the relative ages of marine rocksThere are several resons that fossil foraminifera are especially valuable for determining the relative ages of marine rock layers. They have been around since the Cambrian, over 500 million years ago. They show fairly continuous evolutionary development, so different species are found at different times. Forams are abundant and widespread, being found in all marine environments. Finally, they are small and easy to collect, even from deep oil wells.

PALEOECOLOGY AND PALEOBIOGEOGRAPHY

Foraminifera provide evidence about past environmentsBecause different species of foraminifera are found in different environments, paleontologists can use the fossils to determine environments in the past. Foraminifera have been used to map past distributions of the tropics, locate ancient shorelines, and track global ocean temperature changes during the ice ages. If a sample of fossil foraminifera contains many extant species, the present-day distribution of those species can be used to infer the environment at that site when the fossils were alive. If samples contain all or mostly extinct species, there are still numerous clues that can be used to infer past environments. These include species diversity, the relative numbers of planktonic and benthic species, the ratios of different shell types, and shell chemistry.

The chemistry of the shell is useful because it reflects the chemistry of the water in which it grew. For example, the ratio of stable oxygen isotopes depends on the water temperature, because warmer water tends to evaporate off more of the lighter isotopes. Measurement of stable oxygen isotopes in planktonic and benthic foram shells from hundreds of deep-sea cores worldwide have been used to map past surface and bottom water temperatures. This data helps us understand how climate and ocean currents have changed in the past and may change in the future.

OIL EXPLORATION

Foraminifera are used to find petroleumSome species are geologically short-lived and some forms are only found in specific environments. Therefore, a paleontologist can examine the specimens in a small rock sample like those recovered during the drilling of oil wells and determine the geologic age and environment when the rock formed. As a result, since the 1920's the oil industry has been an important employer of paleontologists who specialize in these microscopic fossils. Stratigraphic control using foraminifera is so precise that these fossils are even used to direct sideways drilling within an oil-bearing horizon to increase well productivity.

BIOLOGY OF FORAMINIFERA

Very little is known about how most species of foraminifera live. The few species that have been studied show a wide range of behaviors, diet, and life cycles. Individuals of some species live only a few weeks, while other species live many years. Some benthic species burrow actively, though slowly, through sediment at speeds up to 1cm per hour, while others attach themselves to the surface of rocks

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or marine plants. Foraminifera are abundant enough to be an important part of the marine food chain, and their predators include marine snails, sand dollars and small fish.

CLASSIFICATION OF FORAMINIFERA

Traditionally, classification of foraminifera has been based primarily on characters of the shell or test. Wall composition and structure, chamber shape and arrangement, the shape and position of any apertures, surface ornamentation, and other morphologic features of the shell are all used to define taxonomic groups of foraminifera. New research is adding molecular data on relationships among species that may greatly affect how these organisms are classified.

Chamber arrangements commonly found in living species are shown in figures 1-6. The following terms are used:

Unilocular refers to a shell made of a single chamber Uniserial refers to chambers added in a single linear series Biserial refers to chambers added in a double linear series Triserial refers to chambers added in a triple linear series Planispiral refers to chambers added in a coil within a single plane like the chambered

nautilus Trochospiral refers to chambers added in a coil that forms a spire like a snail shell Milioline refers to an arrangement where each chamber stretches the full length of the shell

and each successive chamber is placed at an angle of up to 180 degrees from the previous, relative to the central axis of the shell

Arborescent refers to an erect, branching series of tubes. Terms such as planispiral-to-biserial and biserial-to-uniserial are used when the mode of

chamber addition changes during growth.

Of the various kinds of wall composition and microstructure found in foraminifera, three basic types are common among living species. Agglutinated shells may be composed of very small particles cemented together and have a very smooth surface, or may be made of larger particles and have a rough surface. Hyaline shells are made of interlocking microcrystals of CaCO3, and typically have a glassy appearance and pores that penetrate the wall. Porcelaneous shell walls are composed of microscopic rod-shaped crystals of CaCO3. These have a milky, translucent to opaque look and generally lack pores beyond the initial chambers. In some porcelaneous species, small depressions in the surface ornamentation give the appearance of pores. Another type of wall structure, called microgranular, is made of tightly packed equidimensional rounded grains of calcite. This wall type is found in many Paleozoic foraminifera including the fusulinids.

Figures 1-6. These images were captured using the Environmental Scanning Electron Microscope at the UC Museum of Paleontology, Berkeley, CA.

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Return to top

 

As regards the history of life on Earth, the Triassic period was significant for a number of reasons.  This was a time of

transition, in which many old forms of life died out, and new, and sometimes even modern, ones appeared.  The fusilinid

foraminifers, lacy bryozoans, rugose corals and trilobites that had characterized the late Paleozoic all disappeared.

Introduction

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The clade Rhizaria of unicellular eukaryotes was named very recently (Cavalier-Smith, 2002), but has rapidly ingratiated itself as an industry standard.  It contains a large number of mostly amoeboid organisms, including such significant groups as the radiolarians and foraminiferans. 

So far, Rhizaria seems to be supported solely by molecular data – there are no morphological characters unique to the clade. Most are biciliate amoeboflagellates, at least at some point in the life cycle – though many have dispensed with flagella altogether.  Pseudopodia are root-like reticulopodia, filopodia and/or axopodia – not broad lobopodia as in Amoeba.  All of these features can, however, be found in members of other clades.  Nevertheless, the Rhizaria are supported by both rRNA and actin trees (Cavalier-Smith & Chao, 2003; Nikolaev et al . 2004 ), and are probably here to stay.

 Foraminifera

Amoeboid organisms characterised by reticulate, granular pseudopodia (hence the often-seen alternative name Granuloreticulosa). Mostly marine; endosymbiotic algae often present. The majority of Foraminifera produce a test of some form or other – mostly calcareous, but agglutinated or organic in more basal forms. One group of basal agglutinated-test Foraminifera became sessile, and a subgroup of this line took to growing to Brobdignagian proportions – the Xenophyophorea. Pawlowski et al. (2003) . 

Foraminifera, especially the calcareous forms, have a fossil record stretching back to the Cambrian (Lee, 1990), and are especially important biostratigraphically.

The Xenophyophorea are either Foraminifera, or possibly the sister group of Foraminifera.  These bizarre, gigantic protists are commonly several centimeters in diameter and are discussed on their own page.

Text © Christopher Taylor 2004.  CT041218

http://www.ucmp.berkeley.edu/foram/foramintro.html

Introduction to the ForaminiferaForaminifera (forams for short) are single-celled protists with shells. Their shells are also referred to as tests because in some forms the protoplasm covers the exterior of the shell. The shells are commonly divided into chambers which are added during growth, though the simplest forms are open tubes or hollow spheres. Depending on the species, the shell may be made of organic compounds, sand grains and other particles cemented together, or crystalline calcite.

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A typical foram : In the picture about, the dark brown structure is the test, or shell, inside which the foram lives. Radiating from the opening are fine hairlike reticulopodia, which the foram uses to find and capture food.

Fully grown individuals range in size from about 100 micrometers to almost 20 centimeters long. A single individual may have one or many nuclei within its cell. The largest living species have a symbiotic relationship with algae, which they "farm" inside their shells. Other species eat foods ranging from dissolved organic molecules, bacteria, diatoms and other single celled phytoplankton, to small animals such as copepods. They move and catch their food with a network of thin extensions of the cytoplasm called reticulopodia, similar to the pseudopodia of an amoeba, although much more numerous and thinner.

Foraminifera: Fossil Record

The oldest fossil foraminifera, from the Cambrian, are simple agglutinated tubes. Calcareous microgranular and porcellaneous tests evolved in the Carboniferous, and calcareous hyaline tests in the Permian. Over time, each of these groups has evolved many different forms, including large complex tests associated with reefs. These groups of large species became abundant when reef environments were widespread, then suffered major extinction when world climate changed and reefs were decimated. The fusulinids were one such group. They had rice-grain shaped tests and evolved into numerous widespread species during the Permian but went extinct at the end of that period when a worldwide mass extinction also eliminated most other reef dwelling organisms.

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The small size of most foraminifera may make them difficult to see, but it makes them much more useful than larger fossils for applications such as petroleum exploration, because there can be thousands of specimens in the small chips of rock collected when drilling a well. In addition, many species of foraminifera are geologically short-lived, and others are only found in specific environments, so a paleontologist can examine the specimens in a sample and determine the geologic age and environment when the rock formed. As a result, since the 1920's the oil industry has been a major employer of paleontologists who specialize in these microscopic fossils. It is unusual to drill an oil well without a paleontologist onsite to determine when the desired oil-bearing rock layer has been reached.

Foraminifera: Life History and Ecology

Most of the estimated 4,000 living species of forams 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.

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 ocean bottom. Many larger animals (including snails, sand dollars, and fish) eat forams, and some are very selective about which species they eat.

Because 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

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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, data such as species diversity, the relative numbers of planktonic and benthic species (planktic: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 us understand how climate has changed in the past and thus how it may change in the future.

Foraminifera: More on Morphology

Foraminiferan shells, or tests, are built of hollow chambers separated by partitions, with small openings called foramina that connect the chambers (they get their name from these foramina). The final chamber (the last one added) has an opening or openings to the exterior, called the aperture. The living organism fills all the chambers in its shell except for one or two of the most recently constructed chambers. Most species build shells with multiple chambers (multilocular) but some species build shells with only a single chamber (unilocular). Click here to learn more about the most common types of chamber arrangements.

Each of the major groups of foraminifera uses different materials to build their shells. The basic types of wall structures are:

agglutinated -- test made of particles cemented together. Some species use whatever particles are available, while other species may select only sponge spicules or mica flakes or a certain size particle to build their test.

calcareous hyaline -- interlocking crystals of calcite about 1 micrometer in diameter. microgranular -- equidimensional, subspherical particles of calcite closely packed together

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porcellaneous -- wall made of apparently randomly arranged microscopic rods of calcite, with ordered inner and outer surface layers.

Fusulinid limestone from Pakistan. Note the large size of these fusilinids.

Close up of a transverse section of a fusulinid in limestone from Pakistan. Note the internal structure.

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More fusilinid limestone from Pakistan showing fusulinids in every orientation.

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FusulinidFusulinids

Fossil range: Silurian - Permian

Scientific classificationDomain: EukaryotaKingdom: RhizariaPhylum: ForaminiferaOrder: Fusulinida

The fusulinids are an extinct group of foraminiferan protozoa. They produced calcareous shells, which are of fine calcite granules packed closely together; this distinguishes them from other calcareous forams, where the test is usually hyaline. Their fossils are so abundant that they have formed entire limestone formations. The fusulinid Cottonwood Limestone formation in Kansas is an example of this. Fusulinids are important indicator fossils.

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Fusulinids appeared late in the Mississippian Period. They were a part of the Carboniferous and Permian marine communities. They are excellent guide fossils for Pennsylvanian and Permian rocks. However, fusulinids became extinct at the end of the Permian Period.

Fusilinid - tiny football shaped microfossils that have been very well studied because of their association with oil-producing rocks.

Fusilinid (5mm)

This is a 5mm single cell organism! It's approximate age... 250-320 million years old (MYO). Found where there used to be a used to be a Permian sea. Drexel MO/KS. They became extinct during the largest mass extinction on this planet (93-95% all species) the Permian Triassic extinction event. P-

Fossil Fusulinid - Yabeina globosa

Fusulinids are protists from the phylum granuloreticulosa, and the class foraminifera, also called foraminiferida. What does this mean? Granuloretuculosa means that they are cells that fuse to form networks of cells. Foraminifera means it has a "test", or internal skeleton with pores on the outside, through which it feeds with reticulopodia, which are much like the pseudopods of amoebas but are smaller and more hair-like. These protists are common throughout the geological record and are often used as a reference for dating rocks. The fusilinid that I have is a beautiful example of a foraminiferid, or foram for short. Below are several fusulinids in cross-section that have been polished to show their details.

About this specimen

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This is a specimen of several Yabeina globosa from the Upper Permian, Kinsho-zan, Akasaka, Gifu, Japan. Kinsho-zan means Mountain Kinsho, or just Mount Kinsho. Akasaka is the town near where it was found, and Gifu is the district.

(above) Polished section of rock with the fusulinids and a close-up of a fusilinid cross-section. (below) An un-polished section with the fusulinid on the right still partially rounded.

NummuliteNummulites

Fossil range: Tertiary

Scientific classificationKingdom: RhizariaPhylum: ForaminiferaOrder: RotaliidaSuperfamily: Nummulitacea

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Family: NummulitidaeGenus: Nummulites

Lamarck, 1801

Species

Numerous

Fossil nummulitid foraminiferans showing microspheric and megalospheric individuals; Eocene of the United Arab Emirates; scale in mm.

Fossil Nummulites in Urbasa, Basque Country

A nummulite is a large lenticular fossil, characterized by its numerous coils, subdivided by septa into chambers. They are the shells of the fossil and present-day marine protozoan Nummulites, a type of foraminiferan. Nummulites commonly reach 6 cm (2.4 inches) in diameter, and are common in Tertiary marine rocks, particularly around the Mediterranean (e.g. Eocene limestones from Egypt). Fossils up to 6 inches wide are found in the Middle Eocene rocks of Turkey.2 They are valuable as index fossils.

The name "Nummulites" is a diminutive form of the Latin nummulus meaning "little coin", a reference to their shape. In 1913, Randolph Kirkpatrick published a book, The Nummulosphere: an account of the Organic Origin of so-called Igneous Rocks and Abyssal Red Clays, proposing the theory that all rocks have been constructed by the accumulation of forams such as Nummulites.

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nummuliteLarge coin-shaped fossil composed of numerous whorls coiled in a plane spiral, the whorls being divided into chambers by septa. These, the remains of marine protozoa, are common in Tertiary rocks, particularly Eocene limestones in Egypt and elsewhere, and are valuable as index fossils.Nummulites reached the maximum size for any protozoan, of around 6 cm/2.4 in diameter.Classification

Nummulites are in genus Nummulites order Foraminiferida, class Granuloreticulosa, subphylum Sarcodina, phylum Sarcomastigophora

NummuliteNummulites are protists from the phylum granuloreticulosa, and the class foraminifera, also called foraminiferida. What does this mean? Granuloretuculosa means that they are cells that fuse to form networks of cells. Foraminifera means it has a "test", or internal skeleton with pores on the outside, through which it feeds with reticulopodia, which are much like the pseudopods of amoebas but are smaller and more hair-like. Nummulites start out in the center adding a new larger chamber in a spiral pattern, creating a disc-shaped test. Many nummulites can be found in the lime stone of northern Africa and Mediteranean areas. Nummulitic limestone was used in the construction of ancient Egyptian monuments, such as the pyramids. Nummulites were named for the latin word "nummulus", or coin. They were probably named thus because of how it resembles a coin. I have heard that in ancient Egypt, they may have been used as currency, but I cannot confirm this. Some scientists speculate that it may have had algae living inside in a symbiotic relationship.

About this specimen

This is a nummulite from around the Eocene and Oligocene periods 55-34 million years ago, found in the Mediteranean sea in Greece. It's just over an inch at about 30 mm in diameter, which is pretty big for a protist.

Nummulites sp.found at Wemmel/Belgium

Oligocene

Send me your sand or rock ! I extract the foraminifera and shoot the images for free. Info at www.foraminifera.eu/participate. html 

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Nummulitinae miscellaneaRoyan/FranceMaastrichtian

Nummulites sp.found at Wemmel/Belgium

Oligocene

num·mu·lite     (nŭm'yə-līt')  Pronunciation Key  n.   A large, coin-shaped, fossil foraminifer of the genus Nummulites, widely distributed in limestone formations from the Eocene Epoch to the Miocene Epoch of the Cenozoic.

[From New Latin Nummulītēs, type genus, from Latin nummulus, diminutive of nummus, coin, probably from Greek nomimos, customary, legal; see nem- in Indo-European roots.]

http://www.ucl.ac.uk/GeolSci/micropal/foram.html

Foraminifera

Introduction

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Foraminifera are found in all marine environments, they may be planktic or benthic in mode of life. The generally accepted classification of the foraminifera is based on that of Loeblich and Tappan (1964). The Order Foraminiferida (informally foraminifera) belongs to the Kingdom Protista, Subkingdom Protozoa, Phylum Sarcomastigophora, Subphylum Sarcodina, Superclass Rhizopoda, Class Granuloreticulosea. Unpicking this nomenclature tells us that foraminifera are testate (that is possessing a shell), protozoa, (single celled organisms characterised by the absence of tissues and organs), which possess granuloreticulose pseudopodia (these are thread-like extensions of the ectoplasm often including grains or tiny particles of various materials). Bi-directional cytoplasmic flow along these pseudopodia carries granules which may consist of symbiotic dinoflagellates, digestive vacuoles, mitochondria and vacuoles containing waste products; these processes are still not fully understood. In the planktic foraminifera Globigerinoides sacculifer dinoflagellate symbionts are transported out to the distal parts of rhizopodia in the morning and are returned back into the test at night. The name Foraminiferida is derived from the foramen, the connecting hole through the wall (septa) between each chamber.

History of Study

The study of foraminifera has a long history, their first recorded "mention" is in Herodotus (fifth century BC) who noted that the limestone of the Egyptian pyramids contained the large benthic foraminifer Nummulites. In 1835 Dujardin recognised foraminifera as protozoa and shortly afterwards d'Orbigny produced the first classification. The famous 1872 HMS Challenger cruise , the first scientific oceanographic research expedition to sample the ocean floor collected so many samples that several scientists, including foraminiferologists such as H.B. Brady were still working on the material well in to the 1880's. Work on foraminifera continued throughout the 20th century, workers such as Cushman in the U.S.A and Subbotina in the Soviet Union developed the use of foraminifera as biostratigraphic tools. Later in the 20th century Loeblich and Tappan and Bolli carried out much pioneering work.

Range

Foraminifera have a geological range from the earliest Cambrian to the present day. The earliest forms which appear in the fossil record (the allogromiine) have organic test walls or are simple agglutinated tubes. The term "agglutinated" refers to the tests formed from foreign particles "glued" together with a variety of cements. Foraminifera with hard tests are scarce until the Devonian, during which period the fusulinids began to flourish culminating in the complex fusulinid tests of the late Carboniferous and Permian times; the fusulinids died out at the end of the Palaeozoic. The miliolids first appeared in the early Carboniferous, followed in the Mesozoic by the appearance and radiation of the rotalinids and in the Jurassic the textularinids. The earliest forms are all benthic, planktic forms do not appear in the fossil record until the Mid Jurassic in the strata of the northern margin of Tethys and epicontinental basins of Europe. They were probably meroplanktic (planktic only during late stages of their life cycle). The high sea levels and "greenhouse" conditions of the Cretaceous saw a diversification of the planktic foraminifera, and the major extinctions at the end of the Cretaceous included many planktic foraminifera forms. A rapid evolutionary burst occurred during the Palaeocene with the appearance of the planktic globigerinids and globorotalids and also in the Eocene with the large benthic foraminifera of the nummulites, soritids and orbitoids. The orbitoids died out in the Miocene, since which time the large foraminifera have dwindled. Diversity of planktic forms has also generally declined since the end of the Cretaceous with brief increases during the warm climatic periods of the Eocene and Miocene.

Classification

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Foraminifera are classified primarily on the composition and morphology of the test. Three basic wall compositions are recognised, organic (protinaceous mucopolysaccharide i.e. the allogromina), agglutinated and secreted calcium carbonate (or more rarely silica). Agglutinated forms, i.e the Textulariina, may be composed of randomly accumulated grains or grains selected on the basis of specific gravity, shape or size; some forms arrange particular grains in specific parts of the test. Secreted test foraminifera are again subdivided into three major groups, microgranular (i.e. Fusulinina), porcelaneous (i.e. Miliolina) and hyaline (i.e. Globigerinina). Microgranular walled forms (commonly found in the late Palaeozoic) are composed of equidimensional subspherical grains of crystalline calcite. Porcelaneous forms have a wall composed of thin inner and outer veneers enclosing a thick middle layer of crystal laths, they are imperforate and made from high magnesium calcite. The hyaline foraminifera add a new lamella to the entire test each time a new chamber is formed; various types of lamellar wall structure have been recognised, the wall is penetrated by fine pores and hence termed perforate. A few "oddities" are also worth mentioning, the Suborder Spirillinina has a test constructed of an optically single crystal of calcite, the Suborder Silicoloculinina as the name suggests has a test composed of silica. Another group (the Suborder Involutina) have a two chambered test composed of aragonite. The Robertinina also have a test composed of aragonite and the Suborder Carterina is believed to secrete spicules of calcite which are then weakly cemented together to form the test.

The morphology of foraminifera tests varies enormously, but in terms of classification two features are important. Chamber arrangement and aperture style, with many subtle variations around a few basic themes. These basic themes are illustrated in the following two diagrams but it should be remembered that these are only the more common forms and many variations are recognised.

Planktic

Globigerina bulloides d'Orbigny

Pliocene-Recent South Africa380 microns spiral view SEM

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Globigerina bulloides d'Orbigny Pliocene-Recent South Africa416 microns umbilical view SEM

Globigerinoides ruber d'Orbigny Miocene-Recent South Africa

spiral view SEM

Globigerinoides sacculifer (Brady) Miocene-Recent South Africa

spiral view SEM

Globorotalia inflata d'Orbigny Pliocene-Recent South Africaspiral view SEM

Globorotalia inflata d'Orbigny Pliocene-Recent South Africaumbilical view SEM

Globorotalia menardii (Parker, Jones and Brady) Pliocene-Recent South Africa

spiral view SEM

Globorotalia menardii (Parker, Jones and Brady) Pliocene-Recent South Africaumbilical view SEM

Neogloboquadrina pachyderma (Ehrenberg) Pliocene-Recent South Africaspiral view SEM

Neogloboquadrina pachyderma (Ehrenberg) Pliocene-Recent South Africaumbilical view SEM

Orbulina universa d'Orbigny Middle Miocene-Recent South Africa

SEM

Hantkenina alabamensis Cushman, 1927 Eocene

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Montgomery Landing, Red River, Louisiana, USA

side view (slightly broken specimen) SEM

Pseudohastigerina micra (Cole, 1927) Eocene-Oligocene Montgomery Landing, Red River, Louisiana, USA

side view SEM

Globorotalia centralis Cushman and Bermudez, 1937 Eocene Montgomery Landing, Red River, Louisiana, USAumbilical view SEM

Globorotalia cerro-azulensis Cole, 1928 Eocene Montgomery Landing, Red River, Louisiana, USAumbilical view SEM

Parasubbotina varianta (Subbotina, 1953) Lower-Middle Palaeocene Zin Valley, Israel

spiral view SEM

Parasubbotina pseudobulloides (Plummer, 1926)

Lower-Middle Palaeocene Zin Valley, Israel

umbilical view SEM

Subbotina triloculinoides (Plummer, 1926) lower-upper Palaeocene Zin Valley, Israelumbilical view SEM

Subbotina triloculinoides (Plummer, 1926) Palaeocene Zin Valley, Israel

spiral view SEM

Abathomphalus mayaroensis (Bolli) Upper Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syriaumbilical view SEMAbathomphalus mayaroensis (Bolli) Upper Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syrialateral view SEM

Contusotruncana contusa (Cushman) Upper Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syriadorsal view SEM

Contusotruncana contusa (Cushman)

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Upper Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syriaventral view SEM

Globotruncana linneiana (d'Orbigny) Santonian-Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syriadorsal view SEM

Globotruncana linneiana (d'Orbigny) Santonian-Maastrichtian (Upper Cretaceous) Kassbah, N.W. Syriaventral view SEM

Racemiguembelina fructicosa (Egger) Middle-Uppper Maastrichtian (Upper Cretaceous) Kassbah, N.W. SyriaSEM

Racemiguembelina fructicosa (Egger) Middle-Uppper Maastrichtian (Upper Cretaceous) Kassbah, N.W. SyriaSEM

Pseudotextularia elegans (Rzehak) Campanian-Maastrichtian (Upper Cretaceous) Kassbah, N.W. SyriaSEM

Pseudoguembelina excolata (Cushman) Campanian-Maastrichtian (Upper Cretaceous) Kassbah, N.W. SyriaSEM

Archaeoglobigerina cretacea (d'Orbigny) Coniacian-Maastrichtian (Upper Cretaceous) Sens, N. Francescale bar 100 microns edge view SEM

Archaeoglobigerina cretacea (d'Orbigny) Coniacian-Maastrichtian (Upper Cretaceous) Sens, N. Francescale bar 100 microns dorsal view SEM

Archaeoglobigerina cretacea (d'Orbigny) Coniacian-Maastrichtian (Upper Cretaceous) Sens, N. Francescale bar 100 microns ventral view SEM

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Hedbergella delrioensis (Carsey) Coniacian-Santonian (Upper Cretaceous) Faircross, UKscale bar 100 microns ventral view SEM

Whiteinella baltica Douglas and Rankin Coniacian-Santonian (Upper Cretaceous) Winterbourne, UKscale bar 100 microns ventral view SEM

Heterohelix pulchra (Brotzen) Coniacian-Maastrichtian (Upper Cretaceous) N. Norfolk, UKscale bar 100 microns side view SEM

Heterohelix globulosa (Ehrenberg) Coniacian-Maastrichtian (Upper Cretaceous) Sens, N. Francescale bar 100 microns side view SEM

Hedbergella planispira (Tappan) Aptian-Coniacian (Upper Cretaceous) Karai, S.E. Indiaventral view SEM

Hedbergella sigali Moullade Barremian-Aptian (Lower Cretaceous) Karai, S.E. Indiaventral view SEM

Ticinella primula Luterbacher Albian (Lower Cretaceous) Karai, S.E. Indiaventral view SEM

BenthicSpiroloculina ornata (d'Orbigny) -Recent Sea of Marmaraside view SEM

Elphidium macellum (Fichtel and Moll) -Recent Sea of Marmaraside view SEM

Brizalina alata (Seguenza) -Recent Sea of Marmaraside view SEM

Cassidulina neocarinata (Thalmann) -Recent Sea of Marmaraventral view SEM

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Siphotextularia concava (Karrer) -Recent Sea of Marmaraside view SEM

Bigenerina nodosaria (d'Orbigny) ??-Recent Sea of Marmaraside view SEM

Planorbulina mediterranensis (d'Orbigny) ??-Recent Sea of Marmaraunattached side SEM

Nonionella opima (Cushman) ??-Recent Sea of Marmaraside view SEM

Lagena striata (d'Orbigny) ??-Recent Sea of Marmaraside view SEM

Alveovalvulina suteri Bronnimann

Gulf of MexicoLM

Cyclamina cancellata Brady

Gulf of Mexico1000 microns LM

Cyclamina acutidorsata (von Hantken) Oligocene-Miocene Gulf of Mexico1000 microns LM

Cyclamina acutidorsata (von Hantken)

Gulf of Mexico1000 microns LM

Reticulophragmium rotundidorsata (von Hantken) Eocene-Miocene Gulf of Mexico600 microns LM

Cribrostomoides sp.

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Gulf of Mexico600 microns side view LM

Cribrostomoides sp.

Gulf of Mexico600 microns apertural view LM

Haplophragmoides bradyi (Robertson)

Gulf of Mexico600 microns side view LM

Haplophragmoides bradyi (Robertson)

Gulf of Mexico600 microns apertural view LM

Jarvisella karamatensis Bronnimann

Gulf of Mexico600 microns apertural view LM

Recurvoides azuamensis Bermudez Oligocene? Gulf of Mexico1500 microns apertural view LM

Textularia tatumi Cushman and Ellisor Miocene? Gulf of MexicoSEM

Valvulina flexilis Cushman and Renz Oligocene-Miocene Gulf of MexicoSEM

Nodosaria sp. Jurassic Villers sur Mer, Normandy, FranceSEM

Nodosaria sp. Jurassic Villers sur Mer, Normandy, Franceclose up of aperture SEM

Vaginulina bernardi Paalzow Oxfordian (Jurassic) Villers sur Mer, Normandy, France

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SEM

Ammobaculites coprolithiformis Schwager Bathonian-Kimmeridgian (Jurassic) Villers sur Mer, Normandy, FranceSEM

Psammosphaera sp. Schulze middle Ordovician-Recent Voring Basin, offshore NorwaySEM

Miliamina fusca (Bradey) ?-Recent Voring Basin, offshore NorwaySEM

Quinqueloculina impressa Reuss ?-Eocene Voring Basin, offshore NorwaySEM

Larger Benthic

Helicolepidina cf nortoni Vaughan ?Eocene Barinas, S.W VenezuelaThin Section

Heterostegina sp d'Orbigny ?Eocene Barinas, S.W VenezuelaTS

Lepidocyclina tobleri panamensis (Cushman) ?Eocene Barinas, S.W VenezuelaTS

Lepidocyclina pustulosa tobleri (Douville) ?Eocene Barinas, S.W VenezuelaTS

Lepidocyclina pustulosa pustulosa (Douville) ?Eocene Barinas, S.W VenezuelaTS

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Discocyclina (Discocyclina) marginata (Cushman) ?Eocene Barinas, S.W VenezuelaTS

Lepidocyclina cf tobleri (Cushman) ?Eocene Barinas, S.W VenezuelaTS

Nummulites sp. Lamarck Paleocene-Holocene Ainsa, Southern Pyrennees, SpainThin section

Discocyclina sp. Gumbel Mid Paleocene-Upper Eocene Ainsa, Southern Pyrennees, SpainThin section

Alveolina sp. d'Orbigny Upper Paleocene-Upper Eocene Ainsa, Southern Pyrennees, SpainThin section

Alveolina sp. d'Orbigny

Upper Paleocene-Upper Eocene Ainsa, Southern Pyrennees, SpainThin section

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