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UNIVERSITY OF SOUTHAMPTON<br />
ABSTRACT<br />
FACULTY OF SCIENCE<br />
SCHOOL OF OCEAN AND EARTH SCIENCE<br />
Doctor <strong>of</strong> Philosophy<br />
REPRODUCTION AND LARVAL BIOLOGY OF NORTH ATLANTIC ASTEROIDS<br />
RELATED TO THE INVASION OF THE DEEP SEA<br />
by Francisco Benitez Villalobos<br />
A very important objective <strong>of</strong> ecological research is to explain the evolution <strong>of</strong> life histories,<br />
more specifically how natural selection modifies reproduction and development in order to<br />
generate the patterns that are observed in nature. With few exceptions, the reproductive<br />
mechanisms and patterns found in deep-water echinoderms are entirely similar to those found<br />
in shallow-water species. The aims <strong>of</strong> this study were 1) to examine the reproductive biology<br />
<strong>of</strong> the many deep-sea asteroids found on the continental slope to the west <strong>of</strong> Europe in order<br />
to determine if the reproductive adaptations are a function <strong>of</strong> depth, distribution or are<br />
phylogenetically controlled, and 2) to conduct experiments on the effects <strong>of</strong> pressure and<br />
temperature on larval development <strong>of</strong> Atlantic asteroids, to investigate the physiological<br />
potential for deep sea invasion by shallow-water species. Eggs <strong>of</strong> the shallow-water asteroids<br />
Asterias rubens Linnaeus and Marthasterias glacialis (Linnaeus) were fertilized in vitro and<br />
incubated through the early embryonic cleavages until the larval stage. They were subjected<br />
to different temperature/pressure regimes. Early embryos were able to tolerate pressures up to<br />
150 atm at 15 o C and 100 atm at 10 o C. Survivorship <strong>of</strong> A. rubens swimming bipinnaria<br />
remained high (> 70%) after incubation at all the pressure/temperature combinations. In M.<br />
glacialis the highest survival <strong>of</strong> swimming larvae was 100% at 1 atm/5, 15 and 20 o C and 50<br />
atm/15 and 20 o C. Data for the temperature and pressure effects on the later stages <strong>of</strong><br />
development suggest that all the larval stages are more temperature/pressure tolerant than the<br />
early embryos and survivorship becomes greater with larval age. Therefore, the larvae <strong>of</strong><br />
these two species could survive transport to deeper waters and these species may be capable<br />
<strong>of</strong> sending colonists to the deep sea. In the deep NE Atlantic the habitat has selected for<br />
species with specific reproductive traits, which provide them with successful and<br />
advantageous life history strategies. This can be clearly observed in the upper bathyal zone<br />
between 700 and 1100 m, where the environmental conditions have selected for small species<br />
with low fecundity and large eggs, plus habits related directly or indirectly with suspension<br />
feeding. These species exhibit reproductive features with trends to the opportunistic strategy<br />
and are distinctive <strong>of</strong> unpredictable environments, although their large egg size probably<br />
follows the general trend observed in species from cold waters in order to provide the larvae<br />
with energy sufficient for a high survival possibility. Conversely, phylogenetic and<br />
evolutionary factors are also important and seem to be decisive at the deepest waters where<br />
basically mainly species belonging to the strict deep-sea family Porcellanasteridae are found.<br />
All these species possess a mixture <strong>of</strong> features typical <strong>of</strong> classical K strategists and<br />
equilibrium strategists, which enable them to persist in a relatively stable environment with<br />
low energy availability. A comprehensive knowledge <strong>of</strong> the reproductive processes <strong>of</strong> the<br />
deep-sea fauna is essential in order to evaluate the level <strong>of</strong> variability caused in the<br />
environment principally by human activity and the possible effects on life-history <strong>of</strong> the<br />
species.<br />
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