In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits and population differentiation in the insect-pollinated, self-incompatible, perennial herb Primula farinosa. More specifically, I (1) determined whether the effects of floral display and interactions with pollinators and seed predators, and plant reproductive success were frequency-dependent and affected by surrounding vegetation context, (2) examined the consequences of intermittent drought years on population dynamics using numerical simulations based on demographic data collected over seven years, (3) analyzed among-population differentiation in flowering phenology and reproductive allocation, and its relationship to soil-depth at the site of origin. A field experiment suggested that conspicuous plants facilitate inconspicuous plants in terms of pollinator attraction, and that the facilitation effect is contingent on the height of the surrounding vegetation. Further experiments revealed that both mutualistic and antagonistic interactions can result in frequency-dependent selection on floral display. Among inconspicuous plants, both fruit initiation, and damage from seed predators increased with the proportion of the conspicuous morph. The relative strength of these effects, and therefore their net outcome on the relationship between morph ratio and seed production varied among years. I combined information on vital rates and their relation to environmental conditions in simulations to predict future population viability in changing environments. Simulated stochastic population growth rate decreased with increasing frequency of drought years. Reproductive allocation varied significantly among populations both in the field and in a common-garden experiment, but was correlated with soil depth at the site of origin only in the field. The results suggest that among-population variation in reproductive effort in the field mainly reflects plastic responses to environmental conditions, and that this plasticity may be adaptive. The common-garden experiment suggested that the study populations have diverged genetically in flowering time.

<p>In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits and population differentiation in the insect-pollinated, self-incompatible, perennial herb <i>Primula farinosa</i>. More specifically, I (1) determined whether the effects of floral display and interactions with pollinators and seed predators, and plant reproductive success were frequency-dependent and affected by surrounding vegetation context, (2) examined the consequences of intermittent drought years on population dynamics using numerical simulations based on demographic data collected over seven years, (3) analyzed among-population differentiation in flowering phenology and reproductive allocation, and its relationship to soil-depth at the site of origin. </p><p>A field experiment suggested that conspicuous plants facilitate inconspicuous plants in terms of pollinator attraction, and that the facilitation effect is contingent on the height of the surrounding vegetation. Further experiments revealed that both mutualistic and antagonistic interactions can result in frequency-dependent selection on floral display. Among inconspicuous plants, both fruit initiation, and damage from seed predators increased with the proportion of the conspicuous morph. The relative strength of these effects, and therefore their net outcome on the relationship between morph ratio and seed production varied among years. </p><p>I combined information on vital rates and their relation to environmental conditions in simulations to predict future population viability in changing environments. Simulated stochastic population growth rate decreased with increasing frequency of drought years. </p><p>Reproductive allocation varied significantly among populations both in the field and in a common-garden experiment, but was correlated with soil depth at the site of origin only in the field. The results suggest that among-population variation in reproductive effort in the field mainly reflects plastic responses to environmental conditions, and that this plasticity may be adaptive. The common-garden experiment suggested that the study populations have diverged genetically in flowering time.</p>

Soils are the largest pool of carbon (C) in terrestrial ecosystems and store 1500 Gt of C in their soil organic matter (SOM). SOM is a dynamic, complex and heterogeneous mixture, which influences soil quality through a wide range of soil properties. Labile SOM comprises a small fraction of total SOM (approximately 5%), but due to its rapid turnover has been suggested to be most vulnerable to loss following soil disturbance. This research was undertaken to examine the consequences of soil disturbance on labile SOM, its availability and protection in soils using the isotopic analysis of soil-respired CO₂ (δ¹³CO₂). A range of soils were incubated in both the short- (minutes) and long-term (months) to assess changes in labile SOM. Shifts in soil-respired δ¹³CO₂ over the course of soil incubations were found to reflect changes in labile substrate utilisation. There was a rapid depletion of δ¹³CO₂ (from a starting range between -22.5 and -23.9‰, to between -25.8 and -27.5‰) immediately after soil sampling. These initial changes in δ¹³CO₂ indicated an increased availability of labile SOM following the disturbance of coring the soil and starting the incubations. Subsequently δ¹³CO₂ reverted back to the initial, relatively enriched starting values, but this took several months and was due to labile SOM pools becoming exhausted. A subsequent study was undertaken to test if soil-respired δ¹³CO₂ values are a direct function of the amount of labile SOM and soil physical conditions. A range of pasture soils were incubated in the short-term (300 minutes), and changes in soil-respired δ¹³CO₂ were measured along with physical and chemical soil properties. Equilibrium soil-respired δ¹³CO₂, observed after the initial rapid depletion and stabilisation, was a function of the amount of labile SOM (measured as hot water extractable C, HWEC), total soil C and soil protection capacity (measured as specific soil surface area, SSA). An independent experimental approach to assess the effect of SSA, where labile SOM was immobilised onto allophane – a clay mineral with large, active surface area – indicated limited availability of labile SOM through more enriched δ¹³CO₂ (in a range between -20.5 and -20.6 ‰) and a significant (up to three times) reduction in HWEC. In the third study, isotopic measurements were coupled with CO₂ evolution rates to directly test whether equilibrium soil-respired δ¹³CO₂ can reflect labile SOM vulnerability to loss. Soils were sampled from an experimental tillage trial with different management treatments (chemical fallow, arable cropping and permanent pasture) with a range of C inputs and soil disturbance regimes. Soils were incubated in the short- (300 minutes) and long-term (600 days) and changes in δ¹³CO₂ and respiration rates measured. Physical and chemical fractionation methods were used to quantify the amount of labile SOM. Pasture soils were characterised by higher labile SOM estimates (HWEC; sand-sized C; labile C respired during long-term incubations) than the other soils. Long-term absence of plant inputs in fallow soils resulted in a significant depletion of labile SOM (close to 50% based on sand-sized C and HWEC estimates) compared with pasture soils. The values of δ¹³CO₂ became more depleted in 13C from fallow to pasture soils (from -26.3 ‰ to -28.1 ‰) and, when standardised (against the isotopic composition of the solid soil material), Δ¹³CO₂ values also showed a decrease from fallow to pasture soils (from -0.3 ‰ to -1.1 ‰). Moreover, these patterns in isotopic measures were in strong agreement with the amount of labile SOM and its availability across the soils, and were best explained by the isotopic values of the labile HWEC fraction. Collectively, these results confirm that labile SOM availability and utilisation change immediately after soil disturbance. Moreover, isotopic analysis of soil-respired CO₂ is a powerful technique, which enables us to probe mechanisms and examine the consequences of soil disturbance on labile SOM by reflecting its availability and the degree of SOM protection.

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