Alaria esculenta

Alaria (Phaeophyceae, Alariaceae) is a common genus of kelps in the northern hemisphere. Fourteen species are currently recognised of which three (Alaria esculenta (L.) Greville, Alaria pylaii (Bory de Saint-Vincent) Greville, and Alaria grandifolia J. Agardh) are reported for the cold -temperate North Atlantic Ocean. Alaria esculenta, the type species described originally from the North Atlantic, exhibits a range of biogeographically correlated morphotypes suggesting the possibility of multiple specific or intraspecific entities or hybrids (Kraan pers. comm.; Kraan & Guiry 2000 in press). A key to the species of the genus Alaria is given by Widdowson (1971).

• Alaria esculenta forms the main canopy in exposed rocky areas.
• Alaria esculenta is a colonizing species that will occur in recently denuded rock surfaces in exposed to sheltered situations.
• Maximum growth rates in the field (20 cm/month) occur in April and May (Isle of Man). Plants on adjacent rope aquaculture system had an average growth rate of 5 cm per day (Birkett et al., 1999b). Kain & Dawes (1987) reported growth rates of 10 cm per day during one spring growth period on a rope aquaculture system in the Isle of Man. However, fields trials with different strains of Alaria esculenta resulted in highest growth rates of 25 cm/month and lowest rates of 5 cm/month (Kraan pers. comm.).
• In June and July growth slows and the blade becomes eroded and damaged by wave action depending on depth. Alaria esculenta in the lower eulittoral and upper sublittoral will erode away completely due to higher summer water temperatures and bleaching by sunlight. Populations at 2 m or more below low water survive (Kraan pers. comm.).
• In strong currents and low wave action the blade may reach 4 m in length (e.g. Aran Islands, Ireland; Guiry, 1997).
• A short stipe and narrow lamina base is characteristic of exposed conditions whereas in sheltered conditions the stipe is long and the lamina base wider (Widdowson, 1971).

Alaria esculenta is present in the North Pacific and North Atlantic, where it is located north as far as the winter sea ice and as far south as the 16 °C summer isotherm, represented by the French coast of Brittany in the European North Atlantic (Luning, 1990). Its absence in the southern North Sea and English Channel is due to high summer surface temperatures of 16 °C, which it cannot survive (Munda & Luning, 1977; Widdowson, 1971; Sundene, 1962). Its distribution in the Arctic Sea is associated with the -2 °C February winter isotherm (Kraan pers. comm.).

• Dispersal potential varies with phase in the life cycle; zoospores may disperse <2m whereas gametophytes may disperse between 1 -10 m by drifting, maybe up to 100 m (Kraan pers. comm.).
• Alaria esculenta is a perennial and lives for 4 -5 years in the Irish Sea and up to 7 years in Norway.
• Two rows of ligulate sporophylls form in the upper parts of the stipe during spring and in a lesser amount in autumn only (Kraan pers. comm.; Widdowson, 1971).
• New sporophytes appear in early spring and zoospores are produced from sporophylls between October and May (Kraan pers. comm.; Birkett et al., 1998b).
• The sporophylls produce haploid spores (zoospores), by meiosis, that germinate to form the haploid or gametophytic phase (male and female). Gametophytes produce the gametes (sperm and eggs) which fuse after fertilization to form a zygote. The zygotes germinates to form diploid platelets (germlings), the sporophytic phase. Thus the life history is that the large seaweed (sporophyte) alternates with a microscopic filamentous phase (Hoek van den, 1995). Therefore, Alaria esculenta is dioecious and has a heteromorphic diplohaplontic life history.
• The flagellated zoospores are about 5 microns in diameter. They loose their flagella after 24 hrs and settle on the available substrata (Birkett et al., 1998b). However, settling rate is dependant on the local currents, therefore spore settling time is probably longer than 1 day.
• Sundene (1961) noted that next generation sporophytes developed within 10 m of the parent plants in Drobak, Norway. Norton (1992) suggested that the position of the sporophylls close to the substratum may limit dispersal potential, however the local currents are probably of overriding importance for dispersal.
• Gametophytes become fertile in under 10 days in optimal conditions.
• Successful fertilization requires a high density of spore settlement (about 1 mm apart).
• Maturation of the gametophytes can be delayed under less optimal conditions, for example, low light, and development remains vegetative. Fragments of damaged vegetative gametophytes may develop into separate gametophytes (only a few cells are required) hence reproductive potential may be increased. If optimal conditions return the gametophyte may become fertile and produce gametes.
• Spore production may be inhibited by epifauna such as Membranipora membranacea (sea mat) and endophytes such as Streblonema sp.