BIOTIC Species Information for Flustra foliacea
Researched byLizzie Tyler Data supplied byUniversity of Sheffield
Refereed byThis information is not refereed.
General Biology
Growth formTurf
Crustose hard
Feeding methodPassive suspension feeder
Active suspension feeder
Mobility/MovementPermanent attachment
Environmental positionEpibenthic
Typical food typesPhytoplankton, detritus and dissolved organic matter. HabitAttached
BioturbatorNot relevant FlexibilityHigh (>45 degrees)
FragilityFragile SizeMedium(11-20 cm)
HeightUp to 20 cm Growth Rate1.6-3 cm / year
Adult dispersal potentialNone DependencyIndependent
General Biology Additional InformationDetailed diagrams of the autozooid and avicularium of Flustra foliacea are provide by Silén (1977).

Growth form
The newly metamorphosed coronate larvae develops into the first zooid of the new colony, the 'ancestrula'. In its first year of growth, Flustra foliacea forms a flat incrustation on the substratum and commences erect growth during the second year. This is achieved simply by the opposition of actively growing lobes of a colony; on contact two growing edges are deflected vertically (J. Porter, pers. comm.).
The two layers of zooids grow, synchronously 'back to back' forming a bilaminar, erect frond at 90 ° to the original encrusting mat. Branching of the erect fronds varies between branches and colonies (Stebbing, 1971a; Silén, 1981).
Ryland (1976) suggested that erect growth avoids the spatial constraints (availability of substratum and competition) suffered by encrusting forms. Repair of grazing damage, i.e. removal of one bilaminar layer, may result in generation of an new bilateral branch (Stebbing, 1971a).

Growth rates
Stebbing (1971a) reported that growth began in late February/early March but stopped in November in specimens off the Gower Peninsula , with a slight check in growth in August, and no growth occurred over winter. Growth stopped in October in Isle of Man specimens (Eggleston, 1963; cited in Stebbing, 1971a). The winter growth check results in visible annual growth lines, which have been used to age colonies (Stebbing, 1971a; Eggleston, 1972; Menon, 1978). Stebbing (1971a) suggested that the growth line formed a line of weakness, which gave the frond flexibility.

Stebbing (1971a) stated that the length of time spent as an encrusting form was unclear but assumed the first growth line at the base of the frond represented the first winter, 1 years growth. Flustra foliacea colonies regularly reached 6 years of age, although 12 year old specimens were reported off the Gower Peninsula (Stebbing, 1971a; Ryland, 1976). Furthermore, O'Dea & Okamura (2000) demonstrated seasonal fluctuations in zooid size synchronous with temperature regimes, the largest zooid zooids occurring with the lowest temperatures.

Stebbing (1971a) reported that growth rates were reasonably consistent between samples, age classes and years. Stebbing (1971a) reported a mean increment in frond height of 16.8mm/yr, whereas Eggleston (1972) reported that annual lines were usually between 2-3cm apart in Isle of Man specimens, and Menon (1978) reported that Helgoland specimens reached an average of 21.2 mm in height at 2 years old and an average of 79.3 mm after 8 years. Silén (1981) reported that erect fronds grew in zooid number about 10-20 times that of the encrusting base. Menon (1978) reported that growth rates varied in specimens over 5 years old.

At the base of fronds, in the holdfast area, the zooids give rise to layers of non-feeding frontal buds after 3 years of age, which strengthen the base of the frond. The number of layers increases with frond height up to 145mm in height, and up to 20 layers deep (Stebbing, 1971a).

Growth rates probably vary between locations. O'Dea & Okumara (2000) noted that colonies of Flustra foliacea from the Bay of Fundy showed reduced growth compared to colonies in the Menai Straits and the Skagerrak. Low primary productivity, genetic variation and parasitism were cited as possible explanations for the difference.

Regeneration and repair
Silén (1981) reported that experimental removal of a notch in the frond was repaired within 5 -10 days. The newly formed margin grew at normal rates (4-5 zooid lengths per month). Removal of one layer of the bilaminar frond, experimentally (Silén, 1981) or by predators (Stebbing, 1971a) was repaired with similar rapidity, the un-damaged layer, halting growth while the damaged area was repaired (Silén, 1981).

The epizoic fauna of Flustra foliacea was described by Stebbing (1971b) and consisted of 25 species of bryozoan, 5 hydroid species, some sessile polychaetes, barnacles, lamellibranchs and tunicates. The bryozoans Bugula flabellata, Crisia spp. and Scrupocellaria spp. were major epizoites. The stolons of Bugula flabellata penetrate the zooids of Flustra foliacea. Scrupocellaria spp. settled preferentially on the youngest, distal, portions of the frond, possibly to elevate their branches into faster flowing water (Stebbing, 1971b). A small green alga Epicladia flustrae was reported to be a specific epiphyte (Nielsen, 1984). Stebbing (1971a) reported that the growth rates of Flustra foliacea were reduced by ca 50% when encrusted by epizoites. Peters et al, (2003) reported the presence of chemical compounds in Flustra foliacea that demonstrated antagonistic effects against the growth of some associated bacterial species, and electron microscopic examination of the distal end of the zooid revealed no microbial settlement. Dyrynda (1985, cited in Peters et al., 1985) reported the toxicity of extracts of Flustra foliacea on larvae of other modular invertebrates, fish and bacteria.

Some people can react to Flustra sp. and some fishermen have reported allergic reactions to it although this is anecdotal information (J. Porter, pers. comm.). Research into biomedical compounds from marine organisms has revealed that a sample of Flustra foliacea from the southern North Sea yielded deformylflustrabromine, which was moderately cytotoxic to the human colon cancer cell-line HCT-116 (Lysek et al., 2002; Jha & Zi-rong, 2004).
Biology References Hayward & Ryland, 1998, Ryland, 1970, Ryland, 1967, Silén, 1977, Ryland, 1976, Silén, 1981, Stebbing, 1971, Menon, 1978, Stebbing, 1971b, Nielsen R., 1975, Hincks, 1880, Stebbing, 1971a, Jha & Zi-rong, 2004, Lysek et al., 2002, Peters et al., 2003, O'Dea & Okamura, 2000, Hayward & Ryland, 1990, Julie Bremner, unpub data,
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