A biostratigraphic unit is a body of rock defined by its fossil content. It is therefore fundamentally different from a lithostratigraphic unit that is defined by the lithological properties of the rock. The fundamental unit of biostratigraphy is the biozone. Biozones are units of stratigraphy that are defined by the zone fossils (usually species or subspecies) that they contain. In theory they are independent of lithology, although environmental factors often have to be taken into consideration in the definition and interpretation of biozones. In the same way that formations in lithostratigraphy must be defined from a type section, there must also be a type section designated as a stratotype and described for each biozone. They are named from the characteristic or common taxon (or occasionally taxa) that defines the biozone. There are several different ways in which biozones can be designated in terms of the zone fossils that they contain.
Interval biozones These are defined by the occurrences within a succession of one or two taxa. Where the first appearance and the disappearance of a single taxon is used as the definition, this is referred to as a taxon-range biozone. A second type is a concurrent range biozone, which uses two taxa with overlapping ranges, with the base defined by the appearance of one taxon and the top by the disappearance of the second one. A third possibility is a partial range biozone, which is based on two taxa that do not have overlapping ranges: once again, the base is defined by the appearance of one taxon and the top by the disappearance of a second. Where a taxon can be recognised as having followed another and preceding a third as part of a phyletic lineage the biozone defined by this taxon is called a lineage biozone (also called a consecutive range biozone).
Assemblage biozones In this case the biozone is defined by at least three different taxa that may or may not be related. The presence and absence, appearance and disappearance of these taxa are all used to define a stratigraphic interval. Assemblage biozones are used in instances where there are no suitable taxa to define interval biozones and they may represent shorter time periods than those based on one or two taxa.
Acme biozones The abundance of a particular taxon may vary through time, in which case an interval containing a statistically high proportion of this taxon may be used to define a biozone. This approach can be unreliable because the relative abundance is due to local environmental factors. The ideal zone fossil would be an organism that lived in all depositional environments all overthe world and was abundant; it would have easily preserved hard parts and would be part of an evolutionary lineage that frequently developed new, distinct species. Not surprisingly, no such fossil taxon has ever existed and the choice of fossils used in biostratigraphy has been determined by a number of factors that are considered in the following sections.
Rate of speciation
The frequency with which new species evolve and replace former species in the same lineage determines the resolution that can be applied in biostratigraphy. Some organisms seem to have hardly evolved at all: the brachiopod Lingula seems to look exactly the same today as the fossils found in Lower Palaeozoic rocks and hence is of little biostratigraphic value. The groups that appear to display the highest rates of speciation are vertebrates, with mammals, reptiles and fish developing new species every 1 to 3 million years on average. However, the stratigraphic record of vertebrates is poor compared with marine molluscs, which are much more abundant as fossils, but have slower average speciation rates (around 10 million years). There are some groups that appear to have developed new forms regularly and at frequent intervals: new species of ammonites appear to have evolved every million years or so during the Jurassic and Cretaceous and in parts of the Cambrian some trilobite lineages appear to have developed new species at intervals of about a million years. By using more than one species to define them, biozones can commonly be established for time periods of about a million years, with higher resolution possible in certain parts of the stratigraphic record, especially in younger strata.
Depositional environment controls
The conditions vary so much between different depositional environments that no single species, genus or family can be expected to live in all of them. The adaptations required to live in a desert compared with a swamp, or a sandy coastline compared with a deep ocean, demand that the organisms that live in these environments are different. There is a strong environmental control on the distribution of taxa today and it is reasonable to assume that the nature of the environment strongly influenced the distribution of fossil groups as well. Some environments are more favourable to the preservation of body fossils than others: for example, preservation potential is lower on a high-energy beach than in a low-energy lagoon. There is a fundamental problem with correlation between continental and marine environments because very few animals or plants are found in both settings. In the marine environment the most widespread organisms are those that are planktonic (free floating) or animals that are nektonic (free-swimming lifestyle). Those that live on the sea bed, the benthonic or benthic creatures and plants, are normally found only in a certain water depth range and are hence not quite so useful. The rates of sedimentation in different depositional environments are also a factor in the preservation and distribution of stratigraphically useful fossils. Slow sedimentation rates commonly result in poor preservation because the remains of organism are left exposed on the land surface or sea floor where they are subject to biogenic degradation. On the other hand, with a slower rate of accumulation in a setting where organic material has a higher chance of preservation (e.g. in an anoxic environment), the higher concentration of fossils resulting from the reduced sediment supply can make the collecting of biostratigraphically useful material easier. It is also more likely that a first or last appearance datum will be identifiable in a single outcrop section because if sediment accumulation rates are high, hundreds of metres of strata may lie within a single biozone.
Mobility of organisms
The lifestyle of an organism not only determines its distribution in depositional environments, it also affects the rate at which an organism migrates from one area to another. If a new species evolves in one geographical location its value as a zone fossil in a regional or worldwide sense will depend on how quickly it migrates to occupy ecological niches elsewhere. Again, planktonic and nektonic organisms tend to be most useful in biostratigraphy because they move around relatively quickly. Some benthic organisms have a larval stage that is free-swimming and may therefore be spread around oceans relatively quickly. Organisms that do not move much (a sessile lifestyle) generally make poor fossils for biostratigraphic purposes.
Geographical distribution of organisms
Two environments may be almost identical in terms of physical conditions but if they are on opposite sides of the world they may be inhabited by quite different sets of animals and plants. The contrasts are greatest in continental environments where geographical isolation of communities due to tectonic plate movements has resulted in quite different families and orders. The mammal fauna of Australia are a striking example of geographical isolation resulting in the evolution of a group of animals that are quite distinct from animals living in similar environments in Europe or Asia. This geographical isolation of groups of organisms is called provincialism and it also occurs in marine organisms, particularly benthic forms, which cannot easily travel across oceans. Present or past oceans have been sufficiently separate to develop localised communities even though the depositional environments may have been similar. This faunal provincialism makes it necessary to develop different biostratigraphic schemes in different parts of the world.
Abundance and size of fossils
To be useful as a zone fossil a species must be sufficiently abundant to be found readily in sedimentary rocks. It must be possible for the geologist to be able to find representatives of the appropriate taxon without having to spend an undue amount of time looking. There is also a play-off between size and abundance. In general, smaller organisms are more numerous and hence the fossils of small organisms tend to be the most abundant. The problem with very small fossils is that they may be difficult to find and identify. The need for biostratigraphic schemes to be applicable to subsurface data from boreholes has led to an increased use of microfossils, fossils that are too small to be recognised in hand specimen, but which may be abundant and readily identified under the microscope (or electron microscope in some cases). Schemes based on microfossils have been developed in parallel to macrofossil schemes. Although a scheme based on ammonites may work very well in the field, the chances of finding a whole ammonite in the core of a borehole are remote. Microfossils are the only viable material for use in biostratigraphy where drilling does not recover core but only brings up pieces of the lithologies in the drilling mud.
Preservation potential
It is impossible to determine how many species or individuals have lived on Earth through geological time because very few are ever preserved as fossils. The fossil record represents a very small fraction of the biological history of the planet for a variety of reasons. First, some organisms do not possess the hard parts that can survive burial in sediments: we therefore have no idea how many types of worm may have existed in the past. Sites where there is exceptional preservation of the soft parts of fossils (lagerstatten) provide tantalising clues to the diversity of lifeforms that we know next to nothing about. Second, the depositional environment may not be favourable to the preservation of remains: only the most resistant pieces of bone survive in the dry, oxidising setting of deserts and almost all other material is destroyed. All organisms are part of a food chain and this means that their bodies are normally consumed, either by a predator or a scavenger. Preservation is therefore the exception for most animals and plants. Finally, the stratigraphic record is very incomplete, with only a fraction of the environmental niches that have existed preserved in sedimentary rocks. The low preservation potential severely limits the material available for biostratigraphic purposes, restricting it to those taxa that had hard parts and existed in appropriate depositional environments.
