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Establecimiento de cuatro especies de Quercus en el sur de la ...

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Spatial and temporal heterog<strong>en</strong>eity effects on seedling growth and establishm<strong>en</strong>t in four <strong>Quercus</strong> species<br />

the pattern resulting from seed dispersal was suppressed in our case, the <strong>sur</strong>vival<br />

patterns observed may also be influ<strong>en</strong>ced by the spatial structure of the emerg<strong>en</strong>ce.<br />

No spatial aggregation patterns for emerg<strong>en</strong>ce were virtually found. This<br />

suggests that emerg<strong>en</strong>ce is <strong>la</strong>rg<strong>el</strong>y in<strong>de</strong>p<strong>en</strong><strong>de</strong>nt of <strong>en</strong>vironm<strong>en</strong>tal factors or that, as<br />

noted above, the differ<strong>en</strong>tial distribution of <strong>en</strong>vironm<strong>en</strong>tal variables in space conceals<br />

their effect on p<strong>la</strong>nt performance (Gal<strong>la</strong>rdo, 2003; García et al., 2006; Pérez‐Ramos et<br />

al., 2010). The two exceptional emerg<strong>en</strong>ce aggregation patterns correspon<strong>de</strong>d to Q.<br />

ilex in plot 1 and Q. pyr<strong>en</strong>aica in plot 2. The aggregation pattern for Q. ilex was<br />

associated to none of the variables studied. On the other hand, that for Q. pyr<strong>en</strong>aica<br />

was positiv<strong>el</strong>y associated to maximum moisture in spring. Therefore, the least moist<br />

zone in winter has a strong limiting effect on emerg<strong>en</strong>ce of Q. pyr<strong>en</strong>aica seedlings,<br />

consist<strong>en</strong>t with its increased moisture requirem<strong>en</strong>ts (Costa et al., 1997). Success in this<br />

species at the beginning of the dry period was associated to herbaceous biomass,<br />

which facilitated establishm<strong>en</strong>t in those microsites with especially abundant herb<br />

production. Some studies have shown herb species to use <strong>la</strong>rge amounts of water for<br />

grow, thereby limiting <strong>sur</strong>vival of woody p<strong>la</strong>nts at an early growth stage (Davis et al.,<br />

1999; Rey B<strong>en</strong>ayas et al., 2005). However, Q. pyr<strong>en</strong>aica is one of the <strong>Quercus</strong> species<br />

with the highest root investm<strong>en</strong>ts (Quero et al., 2006), which allows it to eva<strong>de</strong> water<br />

competition. In addition, an abundance of herbaceous cover may indicate increased<br />

light and moisture lev<strong>el</strong>s in spring and h<strong>en</strong>ce easier establishm<strong>en</strong>t of oaks at the<br />

emerg<strong>en</strong>ce stage. Although this emerg<strong>en</strong>ce pattern influ<strong>en</strong>ced the <strong>sur</strong>vival pattern at<br />

the beginning of the dry period, p<strong>la</strong>nt mortality increased as drought progressed and<br />

the spatial pattern vanished.<br />

Spatial aggregation in <strong>sur</strong>vival and establishm<strong>en</strong>t success persisted throughout<br />

the studied period. The zones with the lowest <strong>sur</strong>vival rates expan<strong>de</strong>d as the dry<br />

period progressed and the patterns disappeared only wh<strong>en</strong> p<strong>la</strong>nt mortality was<br />

virtually 100% (<strong>de</strong>ciduous species, Q. faginea and Q. pyr<strong>en</strong>aica, in plot 2 at the <strong>en</strong>d of<br />

the summer). Some studies (Quero, 2007a; Herrero et al., 2008) have found spatial<br />

patterns in <strong>sur</strong>vival to differ <strong>de</strong>p<strong>en</strong>ding on whether the climatic conditions were<br />

intermediate (aggregation) or extreme (random mortality). However, these<br />

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