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 strong wave action or tidal
currents such that the sessile habit of many species is advantageous. Firm attachment
prevents the species from being swept away to what might be unfavourable conditions, and
prevents them from being damaged by impact on rocks. In terms of being prostrate or erect
there is a conflict of interest for circalittoral rock species. They are mostly filter
feeders, and they frequently live in vigorous water movement. A prostrate habit protects
them from the worst of the water movement, as well as giving them a very robust
morphology. However, it tends to place them in the boundary layer with limited water
movement, and it is the water which carries their food. Conversely, erect species are more
prone to damage by turbulence or currents and more fully exposed to them; but they are in
a position to maximise food intake. Under conditions of extreme exposure robust
low-growing forms predominate barnacles, massive sponges, short hydroids, bryozoans
and tube-building polychaetes. As exposure moderates the taller erect forms come into
prominence the sea fans, soft corals and the like. These erect forms still tolerate
considerable exposure as they have a tough yet flexible structure which enables them to
withstand turbulence and strong currents without damage.
Recruitment processes
Whilst most 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 typically are not dominated by single species, accepting Alcyonium
digitatum in some biotopes, but support a diverse mosaic of species.
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; 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 succesional 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 10-15 cm in height are between five and ten years old (Hartnoll unpubl.).
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References
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