Heat and drought determine flower female allocation in a hermaphroditic Mediterranean plant family.

In animal-pollinated hermaphroditic species, larger and xenogamous flowers increase male-biased resource allocation, whereas smaller and selfing flowers invest disproportionally more resources to female function. In Cistaceae, an entomophilous and hermaphroditic Mediterranean family, this pattern generally follows a phylogenetic signal. However, resource allocation to carpels is independent of phylogeny, which suggests trait divergences among closely related species during the diversification into different environmental conditions. We tested this hypothesis across 37 species of Cistaceae along a temperature and precipitation gradient, including semiarid, dry, subhumid and humid sites. We quantified the proportions of dry mass and nutrient investment to carpels and tested the influence of the climatic gradient and site-specific precipitation on the interspecific variation in carpel resource allocation. Lowest and highest percentages of resource allocation to carpels ranged from 1.5-4.2% to 24.2-36.6%, respectively. The proportion of resources comprised in carpels significantly decreased with increasing precipitation/decreasing temperature. Thus, carpels comprised proportionally more resources under drier and hotter conditions, especially in semiarid sites. Our results demonstrate how the extent of climatic constraints is more important than phylogenetic relationships in determining stress-induced differences in carpel resource allocation across species of Cistaceae in a Mediterranean environment. We suggest that allocation of proportionally more resources to carpels in drier and hotter sites lies within a strategy to deal with the most stressful conditions by means of a high reproductive effort.

One for all and all for one: retention of colour-unchanged old flowers increases pollinator attraction in a hermaphroditic plant.

Long-lived flowers increase pollen transfer rates, but these entail high water and carbon maintenance costs. The retention of pollinated and reward-free old flowers enhances pollinator visitation to young receptive flowers by increasing floral display size. This mechanism is associated with acropetal inflorescences or changes in flower colour and openness, but the retention of unchanging solitary flowers remains overlooked. We examined pollination-dependent variation in floral longevity and determined stigmatic receptivity, pollen viability and pollen removal rates among flower ages in Kielmeyera regalis, a Neotropical savanna shrub. We also evaluated the effects of floral display size on pollinator visitation rates. Lastly, we determined whether old flowers are unvisited and exclusively increase pollinator attraction to young flowers through flower removal experiments. Regardless of pollination treatment, flowers lasted fully open with no detectable physical changes for 3 days. Over time, stigmas remained receptive but >95% of pollen was removed. Pollinator visitation significantly increased with floral display size and intermediate percentages (15-30%) of newly opened flowers. Accordingly, the retention of reward-free and unvisited old flowers increased young flower-pollinator interaction. Our results reveal the importance of a prolonged floral longevity in increasing pollinator attraction toward newly opened receptive flowers without changes in flower colour and form. We conclude that the retention of pollinated, reward-free and unvisited colour-unchanged old flowers in K. regalis is a strategy that counteracts the water use costs associated with the maintenance of large flowers with increased mate opportunities in a pollen-limited scenario.

Phylogeny determines flower size-dependent sex allocation at flowering in a hermaphroditic family.

In animal-pollinated hermaphroditic plants, optimal floral allocation determines relative investment into sexes, which is ultimately dependent on flower size. Larger flowers disproportionally increase maleness whereas smaller and less rewarding flowers favour female function. Although floral traits are considered strongly conserved, phylogenetic relationships in the interspecific patterns of resource allocation to floral sex remain overlooked. We investigated these patterns in Cistaceae, a hermaphroditic family. We reconstructed phylogenetic relationships among Cistaceae species and quantified phylogenetic signal for flower size, dry mass and nutrient allocation to floral structures in 23 Mediterranean species using Blomberg's K-statistic. Lastly, phylogenetically-controlled correlational and regression analyses were applied to examine flower size-based allometry in resource allocation to floral structures. Sepals received the highest dry mass allocation, followed by petals, whereas sexual structures increased nutrient allocation. Flower size and resource allocation to floral structures, except for carpels, showed a strong phylogenetic signal. Larger-flowered species allometrically allocated more resources to maleness, by increasing allocation to corollas and stamens. Our results suggest a major role of phylogeny in determining interspecific changes in flower size and subsequent floral sex allocation. This implies that flower size balances the male-female function over the evolutionary history of Cistaceae. While allometric resource investment in maleness is inherited across species diversification, allocation to the female function seems a labile trait that varies among closely related species that have diversified into different ecological niches.