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

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108 synthesis<br />

(facilitative) interactions between plant species is considered<br />

separately. For each mechanism, I try to answer the following<br />

questions: Is it theoretically plausible? What is the empirical<br />

evidence that the mechanism is important for diversity maintenance<br />

in real plant communities? I conclude this section with a<br />

discussion <strong>of</strong> open questions.<br />

6.1.1 Equalising mechanisms<br />

As mentioned in chapter 1, equalising mechanisms contribute<br />

to species coexistence by reducing fitness differences between<br />

community members. Numerous trade-<strong>of</strong>fs may equalise the<br />

fitness <strong>of</strong> flowering plants with different strategies, for example,<br />

a trade-<strong>of</strong>f between allocation <strong>of</strong> resources to pollinator<br />

attraction <strong>and</strong> production <strong>of</strong> ovules (Haig & Westoby, 1988),<br />

but these do not affect the fitness disadvantage <strong>of</strong> plant species<br />

with low relative abundance. In the following, I discuss the<br />

effects <strong>of</strong> behavioural traits <strong>of</strong> pollinators <strong>and</strong> the spatial structure<br />

<strong>of</strong> plant communities that may decrease the difference in<br />

the quality <strong>and</strong>/or quantity <strong>of</strong> pollinator visits received by rare<br />

<strong>and</strong> abundant plant species.<br />

Flower constancy, the specialisation <strong>of</strong> individual pollinators<br />

on particular flowering plants, reduces loss <strong>of</strong> pollen to heterospecific<br />

flowers <strong>and</strong> clogging <strong>of</strong> stigmas with pollen <strong>of</strong> other<br />

plant species (Levin & Anderson, 1970; Straw, 1972; Goulson,<br />

1994; Montgomery, 2009). The effect on plant reproduction is<br />

essentially the same as for specialisation at the species level.<br />

However, whether pollinator constancy really reduces the difference<br />

in reproductive success between rare <strong>and</strong> common plant<br />

species depends on the relative numbers <strong>of</strong> pollinators specialising<br />

on each plant species <strong>and</strong> their visitation rates. Obviously,<br />

specialisation on a rare plant species can only be pr<strong>of</strong>itable for<br />

a pollinator if the reward per time it receives is at least as high<br />

as for specialisation on a more common plant species. Since<br />

pollinators foraging on rare plant species have higher costs <strong>of</strong><br />

travelling than those foraging on more abundant plants, the<br />

only way to achieve equal rewards per time is if rare plant<br />

specialists receive a higher reward per flower visited. In a<br />

game-theoretic model <strong>of</strong> pollinator foraging behaviour, Kunin<br />

& Iwasa (1996) explored the conditions for equal pr<strong>of</strong>itability<br />

<strong>of</strong> specialisation on a rare <strong>and</strong> a common plant species, respectively,<br />

<strong>and</strong> the consequences for plant reproductive success. Assuming<br />

that both plant species produce nectar at identical rates,<br />

they found that the rarer plant species always received disproportionately<br />

fewer pollinator visits compared to the common

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