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Those data (Young et al., 1997; Pedrotti, 1990) suggest that such embryos and<br />

larvae could colonise the deep-sea under the warm deep-sea conditions that prevailed<br />

in the late Mesozoic or early Cenozoic (Menzies et al., 1973; Hessler & Wilson,<br />

1983). In the North Atlantic, Tyler & Young (1998) examined the temperature and<br />

pressure tolerances <strong>of</strong> embryos and larvae <strong>of</strong> the shallow water species Echinus<br />

esculentus Linnaeus, shallow and bathyal populations <strong>of</strong> E. acutus Linnaeus and<br />

lower bathyal populations <strong>of</strong> E. affinis (Linnaeus). Embryos and larvae <strong>of</strong> both E.<br />

esculentus and E. acutus were unlike the Mediterranean species, developmental<br />

arrests and abnormalities did not increase with lower temperatures. Embryos <strong>of</strong> E.<br />

acutus var. norvegicus from the bathyal zone tolerated a significantly broader range <strong>of</strong><br />

pressure and temperature than did embryos <strong>of</strong> E. acutus from shallow subtidal habitats<br />

and also developed more rapidly at lower temperatures, suggesting that E. acutus is a<br />

very plastic species and may be currently in the process <strong>of</strong> invading the deep-sea by<br />

slowly adapting to increased pressure.<br />

Source d.f. SS MS F p<br />

Pressure (P) 4 12.693 2.6675 195.23 < 0.001<br />

Temperature (T) 3 14.239 7.7764 552.67 < 0.001<br />

Species (S) 1 1.526 1.971 110.69 < 0.001<br />

P*T 12 6.453 0.0796 56.43 < 0.001<br />

P*S 4 1.63 0.04351 24.12 < 0.001<br />

T*S 3 1.63 0.7820 47.17 < 0.001<br />

P*T*S 12 0.087 0.1132 7.57 < 0.001<br />

Error 80 8.75 0.0134 - -<br />

Table 3.1. Three-way analysis <strong>of</strong> variance for proportion <strong>of</strong> Asterias rubens and<br />

Marthasterias glacialis that developed abnormally after 48 h <strong>of</strong> incubation.<br />

66

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