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the maintenance of species-richness in plant communities

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Ma<strong>in</strong>tenance <strong>of</strong> <strong>species</strong>-<strong>richness</strong> <strong>in</strong> <strong>plant</strong> <strong>communities</strong> 129<br />

1000 r n<br />

I \<br />

I<br />

800 t<br />

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T<br />

I<br />

L<br />

I<br />

I<br />

! 600 I<br />

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I<br />

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f<br />

I<br />

v<br />

I<br />

0 400<br />

I<br />

C<br />

I<br />

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-0<br />

a8<br />

v)<br />

200<br />

0<br />

Fig. 10. The numbers <strong>of</strong> emerged seedl<strong>in</strong>gs <strong>of</strong> Artemisia terrae-albae (-) and Alyssum<br />

desertorurn (---) <strong>in</strong> quadrats <strong>in</strong> <strong>the</strong> Turanian Desert <strong>in</strong> 1965-9 (data from Bykov, 1974).<br />

Pen<strong>in</strong>sula <strong>in</strong> <strong>the</strong> U.S.A. commonly become established on <strong>the</strong> ground (Franklyn &<br />

Dyrness, I 969). Seedl<strong>in</strong>gs <strong>of</strong> <strong>the</strong> dicotyledonous tree We<strong>in</strong>mannia racemosa most <strong>of</strong>ten<br />

become established on <strong>the</strong> trunks <strong>of</strong> tree-ferns <strong>in</strong> <strong>the</strong> warm temperate ra<strong>in</strong>-forest <strong>of</strong><br />

New Zealand (Beveridge, 1973), but this is not true <strong>of</strong> <strong>the</strong> o<strong>the</strong>r <strong>species</strong> <strong>of</strong> tree present.<br />

Published records show that some years favour establishment <strong>of</strong> certa<strong>in</strong> seedl<strong>in</strong>gs<br />

and o<strong>the</strong>r years o<strong>the</strong>r <strong>species</strong> (Daubenmire, 1968a; Bykov, 1974). Thus, <strong>in</strong> one part<br />

<strong>of</strong> <strong>the</strong> Turanian semi-desert, 1952 was good for Artemisia paucyora, Kochia prostrata<br />

and Agropyron pect<strong>in</strong>ifone, 1956 for Tanacetum achillaefolium, Festuca sulcata and<br />

Medicago romanica, 1958 for Astragalus virgatus and Tr<strong>in</strong>ia hispida (Bykov, 1974).<br />

Quantitative records for Artemisia terrae-albae and Alyssum desertmum <strong>in</strong> ano<strong>the</strong>r area<br />

for <strong>the</strong> years 1965-69 are shown <strong>in</strong> Fig. 10. Large differences between years have been<br />

found for heathland <strong>in</strong> Scotland (Miles, 1974) and for forests and meadows <strong>in</strong> F<strong>in</strong>land<br />

(Pertulla, 1941).<br />

The passage <strong>of</strong> <strong>the</strong> young seedl<strong>in</strong>g to <strong>the</strong> sapl<strong>in</strong>g or immature vegetative phase is<br />

only arbitrarily separated from <strong>the</strong> early seedl<strong>in</strong>g phase but most <strong>of</strong> <strong>the</strong> <strong>in</strong>formation<br />

concern<strong>in</strong>g <strong>the</strong> effects <strong>of</strong> <strong>the</strong> physical environment concern <strong>the</strong> later stage. Experiments<br />

have shown that <strong>the</strong> effeets <strong>of</strong> irradiance (NichoIson, 1960; Grime, 1966; Hutch<strong>in</strong>son,<br />

1967) and water supply (Jarvis, 1963; Fenner, 1975) certa<strong>in</strong>ly differentiate <strong>species</strong> <strong>of</strong><br />

<strong>the</strong> same community at this stage <strong>of</strong> <strong>the</strong> life-cycle. The ability to tolerate shade is <strong>of</strong>ten<br />

correlated with large seed-size, at any rate <strong>in</strong> <strong>species</strong> that do not depend on mycorrhiza<br />

for <strong>the</strong>ir organic nutrition as seedl<strong>in</strong>gs (Salisbury, 1941, 1974). However, <strong>the</strong>re<br />

are many exceptions to <strong>the</strong> rule and <strong>the</strong> largest-seeded trees <strong>in</strong> European deciduous<br />

forests (Castanea, Quercm) are all light-demand<strong>in</strong>g (Ellenberg, I 963). Possibly <strong>the</strong>ir<br />

large seeds evolved <strong>in</strong> relation to <strong>the</strong> occupation <strong>of</strong> dry sites and <strong>the</strong> advantage <strong>of</strong><br />

9 BRE 52

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