Association between community assemblage of flower colours and pollinator fauna: a comparison between Japanese and New Zealand alpine plant communities.

BACKGROUND AND AIMS: Flower colour plays a major role in the attraction and decision-making of pollinators. Different functional groups of pollinators tend to prefer different flower colours, and therefor may lead to different flower colour compositions among different communities depending on the visual system of the dominant pollinators. However, few studies have investigated the linkage between pollinator fauna and flower colour composition in natural communities, a theme we explored in the present study. METHODS: Flower spectral reflectance of 106 Japanese and 96 New Zealand alpine plants in the wavelength range 300-700 nm were measured. The composition of pollinator fauna in the communities and the types of pollinators for each plant species were also investigated. KEY RESULTS: Based on bee and fly colour vision models, as well as a principal components analysis, considering phylogenetic non-independence between plant species, flower colours appeared to vary according to pollinator type rather than geographical region. Consequently, flower colour composition differed between the regions, reflecting the bee/fly mixed pollinator fauna of Japan and the fly-dominant pollinator fauna of New Zealand. According to the bee colour vision model, the majority of the colours of hymenopteran-pollinated flowers appeared to be discriminated by bees. In contrast, many of the colours of dipteran-pollinated flowers would not be discriminated by bees and flies. CONCLUSION: The results suggest that the differences in flower colour composition between Japanese and New Zealand alpine communities are due to differences in the pollinator fauna in these communities rather than differences in abiotic factors between the geographical regions and the phylogenetic origin of the communities.

Network modules and hubs in plant-root fungal biomes.

Terrestrial plants host phylogenetically and functionally diverse groups of below-ground microbes, whose community structure controls plant growth/survival in both natural and agricultural ecosystems. Therefore, understanding the processes by which whole root-associated microbiomes are organized is one of the major challenges in ecology and plant science. We here report that diverse root-associated fungi can form highly compartmentalized networks of coexistence within host roots and that the structure of the fungal symbiont communities can be partitioned into semi-discrete types even within a single host plant population. Illumina sequencing of root-associated fungi in a monodominant south beech forest revealed that the network representing symbiont-symbiont co-occurrence patterns was compartmentalized into clear modules, which consisted of diverse functional groups of mycorrhizal and endophytic fungi. Consequently, terminal roots of the plant were colonized by either of the two largest fungal species sets (represented by Oidiodendron or Cenococcum). Thus, species-rich root microbiomes can have alternative community structures, as recently shown in the relationships between human gut microbiome type (i.e., 'enterotype') and host individual health. This study also shows an analytical framework for pinpointing network hubs in symbiont-symbiont networks, leading to the working hypothesis that a small number of microbial species organize the overall root-microbiome dynamics.

Host manipulation of bumble bee queens by Sphaerularia nematodes indirectly affects foraging of non-host workers.

Sphaerularia bombi Dufour is a major parasite of bumble bee queens that manipulates its host's behavior: parasitized queens do not breed and found nests but continue to fly into the early summer months. We examined the indirect consequences of this host manipulation on non-host workers in central Hokkaido Island, Japan. In this area, parasitism of Bombus terrestris by S. bombi is common but does not affect every queen; therefore, as summer begins, B. terrestris queens continue to dominate some flower patches and disappear from others. At sites dominated by parasitized queens, we found that the nectar standing crop of red clover was smaller, B. terrestris workers carried out fewer legitimate visits to red clover and more nectar robberies, and the workers were smaller than at other sites. Removing queens from a site increased the nectar standing crop of red clover, the frequency of worker visits to red clover, and the size of the workers. These results suggest that host manipulation by S. bombi increased competition for flower resources among host queens and non-host workers and altered the interaction between plants and non-host flower visitors.

Inflorescence architecture affects pollinator behaviour and mating success in Spiranthes sinensis (Orchidaceae).

• Despite the wide inflorescence diversity among angiosperms, the effects of inflorescence architecture (three-dimensional flower arrangement) on pollinator behaviour and mating success have not been sufficiently studied in natural plant populations. • Here, we investigated how inflorescence architecture affected inter- and intra-plant pollinator movements and consequent mating success in a field population of Spiranthes sinensis var. amoena (S. sinensis). In this species, the flowers are helically arranged around the stem, and the degree of twisting varies greatly among individuals. The large variation in inflorescence architecture in S. sinensis results from variation in a single structural parameter, the helical angle (the angular distance between neighbour-flower directions). • The numbers of visits per inflorescence and successive probes per visit by leaf-cutting bees decreased with helical angle, indicating that individual flowers of tightly twisted inflorescences received less visitations. As expected from pollinator behaviour, pollinia removal and fruit set of individual flowers decreased with helical angle. Meanwhile, geitonogamy decreased in tightly twisted inflorescences. • Our novel findings demonstrate that natural variation in inflorescence architecture significantly affects pollinator behaviour and reproductive success, suggesting that inflorescence architecture can evolve under pollinator-mediated natural selection in plant populations. We also discuss how diverse inflorescence architectures may have been maintained in S. sinensis populations.

Combined effects of inflorescence architecture, display size, plant density and empty flowers on bumble bee behaviour: experimental study with artificial inflorescences.

Pollen dispersal by pollinators is governed by the extent to which diverse effects on pollinator behaviour act independently or augment or moderate each other. Using artificial inflorescences, we assessed the behavioural responses of bumble bees to inflorescence architecture (raceme, panicle, and umbel), inflorescence size (7 or 13 flowers), inter-inflorescence distance and the proportion of empty flowers per inflorescence. The advantage of large inflorescences in terms of attractiveness was larger for racemes and umbels than for panicles, whereas the effect of inter-inflorescence distance on the number of successive probes was smaller for racemes than for panicles and umbels. The number of flowers probed per visit increased almost proportionally with display size when fewer flowers were empty, whereas the number increased less when many flowers were empty. Our results suggest that display size and the spatial arrangement of flowers and nectar within inflorescences can contribute to efficient pollination by affecting pollinator behaviour interactively.

A test of the effect of floral color change on pollination effectiveness using artificial inflorescences visited by bumblebees.

Floral color change has been recognized as a pollination strategy, but its relative effectiveness has been evaluated insufficiently with respect to other floral traits. In this study, effects of floral color change on the visitation pattern of bumblebees were empirically assessed using artificial flowers. Four inflorescence types were postulated as strategies of flowering behavior: type 1 has no retention of old flowers, resulting in a small display size; type 2 retains old flowers without nectar production; type 3 retains old flowers with nectar; and type 4 retains color-changed old flowers without nectar. Effects of these treatments varied depending on both the total display size (single versus multiple inflorescences) and the pattern of flower-opening. In the single inflorescence experiment, a large floral display due to the retention of old flowers (types 2-4) enhanced pollinator attraction, and the number of flower visits per stay decreased with color change (type 4), suggesting a decrease in geitonogamous pollination. Type-4 plants also reduced the foraging time of bees in comparison with type-2 plants. In the multiple inflorescence experiment, the retention of old flowers did not contribute to pollinator attraction. When flowering occurred sequentially within inflorescences, type-4 plants successfully decreased the number of visits and the foraging time in comparison with type-2 plants. In contrast, floral color change did not influence the number of visits, and it extended the foraging time when flowering occurred simultaneously within inflorescences but the opening of inflorescences progressed sequentially within a plant. Therefore, the effectiveness of floral color change is highly susceptible to the display size and flowering pattern within plants, and this may limit the versatility of the color change strategy in nature.

Temporal variation in floral display size and individual floral sex allocation in racemes of Narthecium asiaticum (Liliaceae).

We analyzed the effects of temporal variation in floral display size (number of flowers open at one time on a plant) on the rate of pollen removed and receipt of individual flowers for the sequential blooming plant Narthecium asiaticum (Liliaceae). Because of the acropetal blooming of this species, the display sizes when upper flowers opened was much greater than the display sizes when lower flowers opened. Our experiments revealed that large displays lead to a high rate of pollen removal from individual flowers, though they do not lead to a high rate of ovule fertilization. Consequently, the rate of pollen grains removed per flower by pollinators was greater in upper flowers than in lower flowers. The pattern of sex allocation in individual flowers within a raceme was consistent with such variation. Namely, both maleness [stamen mass/(stamen mass + pistil mass)] and pollen : ovule ratio were larger in upper flowers. We suggest that the temporal variation in display size, in addition to such factors as dichogamy and pollinator directionality, also produces variation in the probability of successful pollen transfer from individual flowers that may cause the variation in the sex allocation of individual flowers.