The relative contributions of insect and bird pollinators to outcrossing in an African Protea (Proteaceae)

? Premise of the study: A useful, but seldom applied, measure of the effectiveness of different pollinators is their contribution to the rate of outcrossing. This measure is particularly useful in facultatively autogamous plants for which seed set cannot be used as a direct measure of pollinator effectiveness. We used selective exclusion experiments to assess the importance of insects for outcrossing in Protea caffra, a facultatively autogamous shrub with scented flowers that are visited frequently by both birds and insects (mainly beetles). ? Methods and results: Pollen loads on stigmas, pollen tube growth, seed set, seed mass, germination, and early seedling survivorship were similar for vertebrate-excluded and open-pollinated inflorescences. Pollen-supplementation mostly did not increase seed set, revealing resource limitation. Mean multilocus outcrossing rates, estimated using eight polymorphic allozyme loci, were similar for progeny from inflorescences excluded from bird visitors (0.65) and for those visited by both birds and insects (0.59). Wright's fixation indices indicated that the adult population is near Hardy-Weinberg equilibrium but differed markedly for maternal plants (F(IS) = -0.187 ± 0.065) and their early stage progeny (F(IS) = 0.258 ± 0.002). Since seed from self and cross hand-pollinations were equally viable in terms of germination, this discrepancy in F(IS) could be explained by inbreeding depression that occurs between germination and reproductive maturity. ? Conclusions: Since outcrossing rates were not reduced when birds were excluded, we infer that insects are effective agents of cross pollination in P. caffra. This helps to explain the evolution of traits associated with insect pollination, such as fruity floral scent, in this species.     

Variation in scent emission among floral parts and inflorescence developmental stages in beetle-pollinated Protea species (Proteaceae)

Floral fragrances are an important component for pollinator attraction in beetle-pollinated flowers. Several genera in the Proteaceae contain beetle-pollinated species. However, there is no information on the floral scent chemistry of beetle-pollinated members of the family. In this paper we report on the spatial variation and differences between developmental stages in emission of inflorescence (flowerhead) volatiles of four South African Protea species (P. caffra, P. dracomontana, P. simplex, and P. welwitschii) that are pollinated by cetoniine beetles. The scents from different inflorescence parts (bracts, perianth, styles, and nectar) and from successive anthesis stages of whole inflorescences were sampled using dynamic headspace collection and identified using GC–MS. Although the four species shared many scent compounds, possibly reflecting their close phylogenetic relationships and common pollinators, they showed significant differences in overall scent composition due to various species-specific compounds, such as the unique tiglate esters found in the scent of P. welwitschii. The strongest emissions and largest number of volatiles, especially monoterpenes, were from inflorescences at full pollen dehiscence. Senescing inflorescences of two species and nectars of all species emitted proportionally high amounts of acetoin (3-hydroxy-2-butanone) and aromatic alcohols, typical fermentation products. As a consequence, the scent composition of nectar was much more similar among species than was the scent composition of other parts of the inflorescence. These results illustrate how the blends of compounds that make up the overall floral scent are a dynamic consequence of emissions from various plant parts.     

Pollination and breeding system of the enigmatic South African parasitic plant Mystropetalon thomii (Mystropetalaceae): rodents welcome,but not needed

• Unrelated plants adapted to particular pollinator types tend to exhibit convergent evolution in floral traits. However, inferences about likely pollinators from ‘pollination syndromes’ can be problematic due to trait overlap among some syndromes and unusual floral architecture in some lineages. An example is the rare South African parasitic plant Mystropetalon thomii (Mystropetalaceae), which has highly unusual brush‐like inflorescences that exhibit features of both bird and rodent pollination syndromes.
• We used camera traps to record flower visitors, quantified floral spectral reflectance and nectar and scent production, experimentally determined self‐compatibility and breeding system, and studied pollen dispersal using fluorescent dyes.
• The dark‐red inflorescences are usually monoecious, with female flowers maturing before male flowers, but some inflorescences are purely female (gynoecious). Inflorescences were visited intensively by several rodent species that carried large pollen loads, while visits by birds were extremely rare. Rodents prefer male‐ over female‐phase inflorescences, likely because of the male flowers’ higher nectar and scent production. The floral scent contains several compounds known to attract rodents. Despite the obvious pollen transfer by rodents, we found that flowers on both monoecious and gynoecious inflorescences readily set seed in the absence of rodents and even when all flower visitors are excluded.
• Our findings suggest that seed production occurs at least partially through apomixis and that M. thomii is not ecologically dependent on its rodent pollinators. Our study adds another species and family to the growing list of rodent‐pollinated plants, thus contributing to our understanding of the floral traits associated with pollination by non‐flying mammals.

     

Evidence for beetle pollination in the African grassland sugarbushes (Protea: Proteaceae)

Most lineages in the African genus Protea consist of species with large unscented flowers pollinated principally by birds, and several of these lineages also show evidence of shifts to rodent pollination, associated with concealed yeasty-scented flowerheads. In this study we investigated the hypothesis that brightly coloured and fruity-scented flowerheads of four Protea species (P. caffra, P. simplex, P. dracomontana and P. welwitschii) represent a novel shift from bird to insect pollination in a grassland lineage in the genus. These species are visited by a wide range of insects, but cetoniine beetles were found to be the most important pollinators because of their abundance, size and relatively pure pollen loads. Three of the four putatively insect-pollinated Protea species have flowers presented at ground level, and experiments showed that cetoniine beetles preferred inflorescences at ground level to those artificially elevated to the height of shrubs and small trees. Relative to insects, birds were infrequent visitors to all of the study species. The nectar of all the study species contained xylose, as documented previously in bird- and rodent-pollinated Protea species, suggesting that this is a phylogenetically conserved trait. However, the very low concentration of nectar (ca. 8%), short nectar-stigma distance and the fruity scent of florets appear to be traits that are associated with specialisation for pollination by cetoniine beetles.