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1985; Morgan, 1997). These observations support the fact that the acute P. elegans<br />

recruitment phases on Drum Sands were seasonal events rather than responses to non-<br />

seasonally related events.<br />

The timing and duration of recruitment phases have been shown to vary for different<br />

P. elegans populations. The adult population studied by Gudmundsson (1985) at<br />

Cullercoats, North Sea coast of England, were 100% sexually mature during<br />

December and eggs were laid into egg capsules from December to March with<br />

hatching occurring from February to April. A second population studied at the nearby<br />

Blyth Estuary showed a similar reproductive timing, although lagging 1 month behind<br />

the Cullercoats population (Gudmundsson, 1985). Rasmussen (1973) studied P.<br />

elegans populations at Horsen's Fjord and Isefjord. The former population reached<br />

sexual maturity during September, egg-laying occurred between September and March<br />

with larval hatching from November to April. At Isefjord, a shorter reproductive<br />

season was reported with egg-laying not taking place until January and eventuating in<br />

April with larvae hatching between February and May. Muus (1967) found that the<br />

female P. elegans population in the brackish Kysing Fjord had eggs/larvae in their<br />

tubes during only April and May, while Hempel (1957) found that a North Sea<br />

population off the German coast had a more prolonged spawning season, from<br />

February to September.<br />

The differences observed between Kysing Fjord (Muus, 1967) and North Sea<br />

populations (Hempel, 1957) were suggested to have been due to differences in salinity<br />

between the two locations (Muus, 1967). This is supported by the more-or-less<br />

continual recruitment at Drum Sands where the salinity is typically marine with only a<br />

small freshwater input from the River Almond. Additionally, from her observations,<br />

Anger (1984) suggested that reproduction may occur throughout the year in the Baltic<br />

and North Seas, with spring and autumn being the most likely seasons in which<br />

production of offspring mainly takes place. Brey (1991) suggested that a mild winter<br />

led to a prolonged recruitment phase and Muus (1967) stated that higher densities of<br />

P. elegans in the Kysing Fjord occurred as a result of the mild winters of 1956/57.<br />

Therefore, it appears that temperature and salinities are the two main factors<br />

determining the periods of larval recruitment for P. elegans.<br />

76

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