Insects use floral signals to find rewards in flowers, transferring pollen in the process. In unisexual plants, the general view is that staminate (male) and pistillate (female) flowers obtain conspecific pollen transfers by advertising their rewards with similar floral signals. For female plants lacking food rewards, this can lead to floral mimicry and pollination by deceit. In this study, we challenge this view by presenting evidence for different rewards offered by flowers on females and males, as a mechanism promoting sexual dimorphism in Leucadendron xanthoconus (Proteaceae), a clearly sexually dimorphic shrub. The tiny beetle pollinators Pria cinerascens (Nitidulidae) depend entirely on the plants they pollinate for survival and reproduction. Male flowers provide mating and egglaying sites, and food for adults and larvae. Female flowers lack nectar and function to shelter pollinators from rain. Their flower heads have cup-shaped display leaves, and are more closed than are those in males. On rainy days, flowers on females received 30% more visits than did flowers on males, and 90% more than they did on sunny days. When we removed display leaves in females, intact flower heads received 14 times more P. cinerascens visits than did manipulated flower heads, indicating that the cup shape attracts the beetles. In both sexes, having many flowers increased the probability of visits and the number of P. cinerascens visiting a plant. In males, the number of larvae was positively correlated with floral-display size, while in females, seed set (pollen transfers) showed no relationship with floral-display size. Ninety-five per cent of the ovules received pollen and 52% matured into seeds. We explain the sexual dimorphism in L. xanthoconus as a result of an intimate partnership with P. cinerascens pollinators, in conjunction with a rainy climate. Pollinators favour large male floral displays, because they offer a reliable food source for adults and larvae. Frequent rains drive the P. cinerascens to leave males in search of the protection offered by females. Because females offer shelter, an essential resource that is not offered by male plants, they receive sufficient pollen independent of their floral-display size. This pollination system promotes the evolution of sexually dimorphic floral signals, guiding pollinators to different rewards in male and female flowers.

Interspecific interactions can vary within and among populations and geographic locations, and this variation can influence the nature of the interaction (e.g. mutualistic vs. antagonistic) and its evolutionary stability. Globeflowers are exclusively pollinated by flies, whose larvae feed only on their seeds. Here we document geographic variability in costs and benefits in globeflowers in sustaining their pollinating flies throughout the range of this arctic-alpine European plant over several years. A total of 1,710 flower heads from 38 populations were analysed for their carpel, egg and seed contents. Individual and population analyses control for the confounding influences of variation in both: (1) population traits, such as fly density and egg distribution among flower heads; and (2) individuals traits, such as carpel and egg numbers per flower head. Despite considerable variation in ecological conditions and pollinator densities across populations, large proportions (range 33-58%) of seeds were released after predation, with a benefit-to-cost ratio of 3, indicating that the mutualism is stable over the whole globeflower geographical range. The stability of the mutualistic interaction relies on density-dependent competition among larvae co-developing in a flower head. This competition is revealed by a sharp decrease in the number of seeds eaten per larva with increasing larval number, and is intensified by non-uniform egg distribution among globeflowers within a population. Carpel number is highly variable across globeflowers (range 10-69), and flies lay more eggs in large flowers. Most plants within a population contribute to the rearing of pollinators, but the costs are greater for some than for others. Large globeflowers lose more seed to pollinator larvae, but also release more seed than smaller plants. The apparent alignment of interests between fly and plants (positive relationship between numbers of seed released and destroyed) is shown to hide a conflict of interest found when flower size is controlled for.

The balance of intimate relationships between plants and seed-eating pollinators can depend on pollinator behaviour in relation to floral characters, such as flower size and flower number. Here, we examined how pollinator oviposition in relation to these traits affected annual fitness (seed output) of single- and multi-flowered Trollius europaeus along altitudinal gradients in subarctic Sweden and the French Alps. Small flies (Chiastocheta spp.) pollinate T. europaeus and their larvae feed on developing seeds. Assuming that late flowers in multi-flowered plants attracted flies to the earliest flower on the same plant, we expected more eggs and higher seed predation in early multiple flowers than in single flowers. Relative seed predation would thereby increase with flower number. Both in Sweden and the Alps, more eggs were placed on large flowers. Early multiple flowers were slightly larger than single flowers, and about twice the size of secondary flowers. As a result, and possibly combined with the effects of secondary flowers, early multiple flowers attracted more ovipositing flies and experienced relatively higher seed predation. However, this did not generally result in higher seed predation of multi-flowered hosts. Multiple flowers had greater seed output than single flowers at all altitudes, also in the high alpine and subarctic sites, where single flowers were more abundant. We hypothesise that the distribution of multiple flowers generally is enforced by environmental factors, rather than by fly-host plant interactions, because only very rarely (in triple-flowered alpine plants) was seed predation disproportionate, and the relationship skewed to the disadvantage of the host. The outcome of the mutualistic interaction was often similar in alpine and subarctic populations, but the underlying factors were different. Subarctic flowers had high abortion and low predation rates, while alpine flowers experienced the reversed situation. The higher fly abundance in the Alps suggests a more intense mutualistic interaction than in Sweden. Despite varying ecological and environmental conditions at these sites, the mutualistic relationship was generally in balance. However, when it was unbalanced, this could be explained by fly behaviour in response to floral traits, and by differences in fly abundance.

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Reproductive and somatic biomass, nitrogen (N), and phosphorus (P) pools were compared between females and males in 1st-year plants of Silene dioica. We estimated irretrievable resources allocated to seeds, pollen, flowers, and unrecovered summer leaf investment by collecting plant parts at abscission throughout the season. At the end of the season, we determined resources lost through senescent stems and autumn leaf turnover and resources stored in perennial roots and overwintering buds. Sexual differences in allocation patterns depended on the resource used for comparison, and whether absolute or proportional resource pools were assessed. Total resource pools in terms of biomass and N were similar for females and males. However, male plants acquired relatively more P. The proportional reproductive investment, i.e., reproductive effort, was similar for males and females in terms of biomass and N. In terms of P, male reproductive effort was higher. There was no difference between sexes in the proportional and relative biomass allocated to perennial roots and overwintering buds. However, in terms of absolute and relative N allocation to below-ground parts, females had larger reserves than males. Females, moreover, had a larger proportion of their P in below-ground parts. However, as male total P pools were larger, absolute P reserves did not differ between sexes. The high reproductive effort and N depletion of below-ground parts in males resulted largely from higher flower production compared to females. In females, seeds were the major component of reproductive effort. These results show that if biomass and nutrient allocation are assessed in parallel for dioecious plants, we obtain a more complete view of their sexual differences.