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Linking Specialisation and Stability of Plant ... - OPUS Würzburg

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6.1 mechanisms <strong>of</strong> diversity maintenance 111<br />

(Eckhart et al., 2006). Yet even if this mechanism should be<br />

widespread, the fact that only relatively inefficient or infrequent<br />

pollinators are expected to exhibit negative frequency-dependence<br />

reinforces the competitive disadvantage <strong>of</strong> rare plant species.<br />

Thus, it seems highly unlikely that negative frequencydependence<br />

<strong>of</strong> pollinators can account for diversity maintenance<br />

in animal-pollinated plant communities.<br />

In addition to the potential equalising effect <strong>of</strong> small-scale<br />

spatial structure <strong>of</strong> plant communities, spatial structure at a<br />

larger scale may act as a stabilising mechanism. Whereas in<br />

our model <strong>of</strong> plant-pollinator systems (chapter 2) in the absence<br />

<strong>of</strong> niche differentiation slight disturbances lead to competitive<br />

exclusion <strong>of</strong> all but the most abundant plant species in<br />

a given location, different species may be favoured by chance<br />

in different local communities. In this manner, diversity could<br />

be maintained at a regional scale. Moreover, if dispersal between<br />

local populations is sufficiently strong, declining populations<br />

may be rescued by immigration from patches where<br />

the species dominate (“mass effect”: Leibold et al., 2004, <strong>and</strong><br />

references therein). Since a mass effect results in a higher percapita<br />

“birth rate” (i.e., births plus immigrants) when a species<br />

becomes rare, it fulfils the criterion for a stabilising mechanism.<br />

How important this mechanism is for diversity maintenance<br />

in real plant communities is currently uncertain, due to a lack<br />

<strong>of</strong> both data <strong>and</strong> models. Mutualistic metacommunities have<br />

been modelled by Prakash & de Roos (2004) <strong>and</strong> Fortuna &<br />

Bascompte (2006), but these studies did not account for competitive<br />

interactions within plant <strong>and</strong> animal communities. In<br />

general, diversity can be maintained through a mass effect if<br />

local communities are sufficiently separated to have diverging<br />

dynamics, but sufficiently connected by dispersal to prevent<br />

competitive exclusion <strong>of</strong> declining species. The classical mass<br />

effect is based on spatial heterogeneity <strong>of</strong> the environment that<br />

favours different species in different locations (Leibold et al.,<br />

2004), but in the case <strong>of</strong> animal-pollinated plants dominance<br />

<strong>of</strong> one species in a given location may be the result <strong>of</strong> incidental<br />

higher initial abundance, which would then be amplified<br />

by its higher pollination success. Further research is needed to<br />

assess the importance <strong>of</strong> mass effects for coexistence <strong>of</strong> animalpollinated<br />

plants, but at least for plant species that are generally<br />

rare alternative mechanisms need to be considered.<br />

Another potential stabilising mechanism also involves structured<br />

plant populations, but here the structure is temporal rather<br />

than spatial. Clearly, temporal segregation <strong>of</strong> flowering phenologies<br />

reduces interspecific competition for pollination. In

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