Calcareous nannofossils include the coccoliths and coccospheres of haptophyte algae and the associated nannoliths which are

of unknown provenance. The organism which creates the coccosphere is called a coccolithophore, they are phytoplankton (autotrophs that contain chloroplasts and photosynthesise). Their calcareous skeletons are found in marine deposits often in vast numbers, sometimes making up the major component of a particular rock, such as the chalk of England. One freshwater species has been reported. Formally coccolithophores are separated from other phytoplankton such as diatoms by the presence of a third flagellalike appendage called a haptonema, although the flagella bearing stage is often only one of a multi-stage life cycle.

 A coccolith is a single disc-like plate which is secreted by the algal organism and held in combination with several other, sometimes varying shaped plates by an organic coating to form the coccosphere. On death the individual coccoliths invariably become separated and it is these that are most commonly preserved in the sedimentary record. Occasionally complete coccospheres are preserved and provide valuable information, particularly regarding coccospheres which possess two or more morphologicaly different coccoliths. There are two forms of coccoliths, the holococcoliths which are formed from calcite crystals which are essentially identical in shape and size and the heterococcoliths which are formed from larger calcite crystals which vary in size and shape. Most living forms are known to produce only heterococcoliths and then only during the non-motile stage of their life cycle. Those that do produce holococcoliths do so only during their motile stage.

 First recorded occurrences of calcareous nannofossils (nannoliths) are from the late Triassic (Carnian).
The locations from which the earliest nannofossils are found include; the Northern and Southern Calcareous Alps, Timor, North-West Australia and Queen Charlotte Islands (Canada), all low latitude sites at the time.

There are many claims for earlier occurrences but a lack of substantiated evidence means these must be excluded. One consequence of the first occurrence of calcareous nannofossils in the late Triassic lies in the fact that this was the first time open ocean planktonic organisms utilised calcareous skeletons and exported calcium carbonate into the deep oceans.

 This has important repercussions in terms of biogeochemical cycles. Today coccolithophores are one of the most important forms of phytoplankton found in the oceans, and may be described as the grass of the sea.

 Culture techniques have resulted in great advances in the study of coccolithophore life cycles. The existence of a haploid and diploid phase has been proved by the extraction of DNA, with mitotic reproduction occurring in both stages. Syngamy (sexual reproduction) has not been observed but is assumed to occur, the recent discovery of combination coccospheres (where coccoliths of two distinct forms occur on the same coccosphere) has meant the traditional classification will have to be radically revised and updated.

 The defining feature of the haptophytes is the flagellalike haptonema which is generally coiled. It differs from the flagella proper in its internal structure and its basal attachment. During the nonmotile phase the flagella disappear but the haptonema often remains, the exact function of the haptonema is not fully understood. The algal cell contains a nucleus and two goldenbrown chloroplasts which may be moved around the cell to optimise collection of available light. The cell also contains mitochondria which contain enzymes which produce the energy for cell function, vacuoles which deal with waste products and the Golgi body which is the site of coccolith secretion in many species.

 In many species overlapping oval organic scales coat the outer cell membrane. These have concentric ridges on their distal faces and radiating ridges on their proximal faces. It seems the organic scales act as bases for the precipitation of the calcite coccoliths. A variety of coccolith secretion strategies have been observed in different species, however it is probably true of all coccolithophores that the production of coccoliths is controlled by light. Emiliania huxleyi has been observed to start coccolith production within half an hour of being introduced to light, and produce an individual coccolith in one hour and a complete coccosphere in about thirty hours.

Coccoliths morphology
Plate (coccolith) consists of shields, plates, central area with or without bars, spine, distal side (outward) and proximal side

Morphological components of coccoliths

 The function of coccoliths is not known but may be one or more of four basic possibilities:

Protection; from bacteria, physical damage, predators such as copepods or to form a chemical buffer zone.
Flotation and buoyancy; aspherical forms may reduce sinking rates, the loss or addition of coccoliths may be a strategy employed to regulate position in the water column in order to optimise light or nutrient availability.

Light regulation; coccoliths may reflect sunlight protecting the cell from high light levels in the upper water column or refract sunlight into the cell allowing life in the lower photic zone. Biochemistry; the cell may secrete calcite in order to expell a metabolic by-product enhancing the bichemical efficiency of the cell.

Kingdom: Protista Division: Chrysophyta Class: Coccolithophyceae

Classification is complicated by the fact that some species are dimorphic, that is they possess more than one coccolith on a single coccosphere. This may lead to the belief that two species exist where in fact there is only one.
Also, pleomorphism (where a holococcolith phase alternates with a heterococcolith phase) may also result in coccoliths being placed in different species or even genera when in fact they are simply different stages in the life cycle of the same species.

Coccolith Life-Style, Ecology and Reproduction

Coccolithophores live in the photic zone (the surface waters, where sunlight reaches) and are photosynthesising (autotrophic); so are at the bottom of the marine food chain, excellent food for herbivorous bacteria. Some have flagella (whip-like appendages) so unlike plants, are capable of movement; furthermore, they don't simply float around, but can swim. Although they are photosynthesising, some have been known to eat

bacteria.
Reproduction is asexual-----simple division. Sensitive for temprature, more abundant at the tropics 100 000 cells/l.

Coccoliths and Sedimentation

After death, they sink by rate of 15 cm/day. Factors of dissolution They form high proportion of carbonate in the sediments Recent 26% Chalk (Cretaceous) 26% Eocene 90%
They are largest producers of calcium carbonate on Earth today, and probably have been since the Late Jurassic. They dump about 1.5 million tons yearly of limestone to the ocean floor.

Geologic history of coccoliths

Abundance of coccoliths in the stratigraphic column (Brasier, 1980).

As the groups name suggests calcareous nannofossils are small, generally less than 30 microns across and usually between 5 and 10 microns (individual coccoliths). This has advantages and disadvantages. Advantages include:

Good preservation, their small size makes mechanical damage unlikely.

Widespread distribution, as part of the phytoplankton coccolithophores are distributed throughout the photic zone (predominantly the upper 50m of the water column) across almost all marine habitats.

A very large number of individual coccoliths may be preserved in a tiny amount of sediment hence only very small quantities of sample are needed to produce statistically valid results. Disadvantages include: Because of dissolution of calcium carbonate at depth in sea water (called the carbonate compensation depth (CCD)), preservation is compromised in deep water sediments. Because of their small size and resistance to mechanical breakdown nannofossils can be reworked, great care is therefore needed especially when utilising nannofossils for biostratigraphic studies. Again, because of the small size the opportunities for contamination are high, although careful and thorough preparation and collection techniques should significantly reduce this risk.