Ecological relationships
Environmental factors determine which circalittoral rock species can inhabit a given
location and this will depend on the biological characteristics of each species
such as size, habit, feeding method and reproductive mode. However, not every species
occurs throughout its potential range its realised distribution is moderated by
biological interactions such as competition, grazing and predation. Circalittoral
communities have been poorly studied in respect to biological interactions because in
order to determine the real role of species in a community experimental manipulation in
the field is required. This has been carried out extensively in the intertidal and
infralittoral but rarely in the circalittoral owing to the logistics of working at such
depths.
Habitat complexity
Circalittoral rock provides a firm attachment in areas of moderate wave action or tidal
currents such that the sessile habit of many species may be advantageous. Firm attachment
will prevent the species from being swept away to what might be unfavourable conditions,
and will prevent them from being damaged by impact on rocks. Under conditions of moderate
exposure the taller erect forms such as sea fans, soft corals and the like are prominent.
These species still tolerate considerable exposure though they have a tough yet flexible
structure, which enables them to withstand turbulence and strong currents without damage
and also allows individuals to maximise food intake.
Recruitment processes
Whilst most moderately exposed circalittoral rock species spend their larval life in
the plankton, there are a few planktonic species which spend their early stages within the
circalittoral rock biotopes. This is true, in rather different degrees, of the hydroids
and the jellyfish. Hydroids are common and conspicuous members of circalittoral rock
biotopes, but the attached hydroids are only the juvenile stages. The sexually reproducing
mature stages are small medusae, which are released into the plankton, where they produce
larvae, which settle again. In contrast, for jellyfish the large adult medusae in the
plankton are the prominent phase. The juvenile stages live attached to rocks as an
inconspicuous scyphistoma stage in which the jellyfish overwinters. In spring this buds
off a series of juvenile medusae, or ephyrae, which grow rapidly in the plankton to form
the adult.
Productivity
Although not primary producers circalittoral rock communities are important
secondary producers. They accumulate and concentrate the primary production from a large
water mass, and make this readily available to higher trophic levels.
Keystone (structuring) species
Circalittoral rock biotopes in moderately exposed conditions typically are not
dominated by single species, but support a diverse mosaic of species. The biotopes where
single species dominate (e.g. Modiolus, Sabellaria and brittlestars) are
described separately.
Importance of habitat for other species
Circalittoral rock communities interact with others by the provision of food and /or
temporary shelter to mobile species which are not permanent faunal turf fauna. Shelter is
important to juvenile fish, which can find refuge (and food) amongst the dense turf of
sessile species. A food source is provided to large mobile crustaceans and fish, which are
attracted by the rich and stationary food supply available on circalittoral rock.
Temporal changes
One of the features of circalittoral faunal turf communities is their fine-scale
spatial variation, which tends to be very patchy. Whilst the infralittoral tends to be
more predictable, circalittoral rock tends to be a mosaic of different species patches
such mosaicing is particularly pronounced in moderately exposed circalittoral communities
compared with exposed and sheltered communities. The different assemblages may represent
alternate stable states (Sutherland 1974; Sebens 1985a, b). In most of these
biotopes substratum space is very fully occupied and the availability of space is a
controlling resource for the settlement and growth of species. According to when free
space is made available, and on which species are recruiting at that time, different
assemblages of species may develop under the same physio-chemical conditions. Once
established, often following a successional sequence (Hextall 1994), these assemblages are
stable for long periods and different assemblages may co-exist in close proximity.
Time for community to reach maturity
Information is restricted, but it is clear that a number of the more prominent members
of the circalittoral rock communities are relatively long lived, and fairly slow growing,
some with life spans ranging from 6-100 years. The soft coral Alcyonium digitatum
is a very prominent member of the circalittoral rock community and observations have shown
that colonies of 10-15cm in height are between five and ten years old (Hartnoll unpubl.).
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References
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