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Revue internationale d'écologie méditerranéenne International ...

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INTRODUCTION<br />

ecologia mediterranea, tome 29, fascicule 2, 2003, p. 153-164<br />

INTERPRETING GERMINATION RESULTS BASED ON SEED SIZE, MASS AND ECOLOGICAL… ◆<br />

Most studies on the evolution or ecological significance<br />

of seed size begin with two observations. Firstly,<br />

across the global flora, seed size varies over some 10<br />

orders of magnitude, and secondly, within species, seed<br />

size is remarkably constant (Harper et al., 1970; Silvertown,<br />

1989). An explanation for the apparent constancy<br />

within species was provided by the theoretical treatment<br />

of Smith and Fretwell (1974), who assumed that a plant<br />

has fixed amount of resources to allocate for reproduction<br />

(reproductive efforts) and that a decision must therefore<br />

be made concerning both the number and size of the<br />

offspring.<br />

Differences in seed size among species may represent<br />

an adaptation to different types of micro-sites<br />

(Silvertown, 1989). If species compete for establishment<br />

sites, large seeded species must have some advantage<br />

to counterbalance the reduction in their seed production<br />

(Lindsay et al., 1999). This advantage could take a<br />

number of forms: (i) small-seeded species could suffer<br />

higher levels of predation, (ii) large-seeded species being<br />

the best competitor for all sites, (iii) safe-micro-sites<br />

for the large-seeded species being much more frequent<br />

than those for small-seeded species. The latter form is<br />

in agreement with numerous experimental studies that<br />

have shown that larger-seeded species are better able to<br />

establish and survive under a wide range of conditions<br />

(Westoby et al., 1997).<br />

There are a number of arguments as to why large<br />

seeds should be advantageous in low moisture conditions.<br />

Baker (1972) and Salisbury (1975) argued that the large<br />

seed weight may enable the seedling to allocate proportionately<br />

more to root development and so produce<br />

extensive root system quickly. For example, in Central<br />

Australia seedlings issued from larger seeds had larger<br />

roots up to 10 days after germination than seedlings from<br />

smaller seeds of 32 species (Jurado & Westoby, 1992).<br />

Several studies have shown that intra-specific mortality<br />

rates in the field are higher for smaller seedlings under<br />

water stress (Parker, 1982; Cook, 1980). Thus, the larger<br />

root systems of seedlings from large seeds may provide<br />

an advantage by allowing access to soil moisture at<br />

deeper levels. The final argument concerning the likely<br />

advantage of large seed size concerns the energetic cost<br />

of tolerating low moisture conditions. Large seeds provide<br />

a larger metabolic reserve for seedlings than small seeds.<br />

Morphological and physiological characteristics, which<br />

confirm drought tolerance, are energetically expensive, so<br />

that the probability of establishment in low soil moisture<br />

155

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