Diel fragrance pattern correlates with olfactory preferences of diurnal and nocturnal flower visitors in Salix caprea (Salicaceae)

Floral scent, often a complex mixture of several volatile organic compounds (VOCs), has generally been interpreted as an adaptation to attract pollinators. However, not many studies have analysed which VOCs are functionally relevant for the reproductive success of a plant. Here, we show that, in Salix caprea (Salicaceae), temporal changes in floral scent emission during the day and night attract two different types of flower visitor: bees during the day and moths during the evening and night. We analysed the contribution of the two flower visitor groups to the reproductive success of the plant. The differences in scent emitted during the peak activity times of flower visitors (day versus night) were quantified and the response of 13 diurnal/nocturnal pollinator taxa to the floral scents was tested using gas chromatographic and electroantennographic techniques. Many of the c. 40 identified scent compounds were physiologically active, and bees and moths responded to nearly identical sets of compounds, although the response strengths differed. In bioassays, bees preferred the most abundant 1,4‐dimethoxybenzene over lilac aldehyde, a compound with increased emission at night, whereas moths preferred lilac aldehyde over 1,4‐dimethoxybenzene. Pollination by wind plus nocturnal pollinators (mainly moths) or by wind alone contributed less to seed set than pollination by wind plus diurnal pollinators (mainly bees). This suggests that the emission of scent during the night and attracting moths have no significant effect on reproductive success. It is possible that the emission of lilac aldehydes and other compounds at night is s result of phylogenetic constraints. Future studies should investigate whether moths may produce a marginal fitness gain in some years and/or some populations. © 2014 The Linnean Society of London, Botanical Journal of the Linnean Society, 2014, 175 , 624–640.     

Consequences of floral visits by ants and invasive honeybees to the hummingbird‐pollinated,Caribbean cactus Melocactus intortus

The pink, tubular, nectariferous flowers of Melocactus intortus (Cactaceae) in Puerto Rico are visited by native hummingbirds (Anthracothorax dominicus), but also by invasive honeybees (Apis mellifera) and ants (Solenopsis sp.). We sought to determine if the bees and ants significantly alter the pollination of M. intortus by measuring pollinator effectiveness. Using traditional estimates of effectiveness (visitation rate and seed set), our results show that hummingbirds were the most effective pollinators as expected. Bees and ants were less effective, and their contributions were one‐fourth to one‐tenth of that observed for hummingbirds. We then modified this measure of effectiveness by adding two components, fitness of progeny and temporal availability of visitors, both of which refine estimates of flower visitor effectiveness. With these new estimations, we found that the effectiveness values of all three animal visitors decreased; however, the role of hummingbirds as the principal pollinator was maintained, whereas the effectiveness values of bees and ants approached zero. By these new measures of overall pollinator effectiveness, the invasive honeybees and ants have little effect on the reproductive success of M. intortus.     

Pollinator genetics and pollination: do honey bee colonies selected for pollen‐hoarding field better pollinators of cranberry Vaccinium macrocarpon?

1. Genetic polymorphisms of flowering plants can influence pollinator foraging but it is not known whether heritable foraging polymorphisms of pollinators influence their pollination efficacies. Honey bees Apis mellifera L. visit cranberry flowers for nectar but rarely for pollen when alternative preferred flowers grow nearby. 2. Cranberry flowers visited once by pollen‐foraging honey bees received four‐fold more stigmatic pollen than flowers visited by mere nectar‐foragers (excluding nectar thieves). Manual greenhouse pollinations with fixed numbers of pollen tetrads (0, 2, 4, 8, 16, 32) achieved maximal fruit set with just eight pollen tetrads. Pollen‐foraging honey bees yielded a calculated 63% more berries than equal numbers of non‐thieving nectar‐foragers, even though both classes of forager made stigmatic contact. 3. Colonies headed by queens of a pollen‐hoarding genotype fielded significantly more pollen‐foraging trips than standard commercial genotypes, as did hives fitted with permanently engaged pollen traps or colonies containing more larvae. Pollen‐hoarding colonies together brought back twice as many cranberry pollen loads as control colonies, which was marginally significant despite marked daily variation in the proportion of collected pollen that was cranberry. 4. Caloric supplementation of matched, paired colonies failed to enhance pollen foraging despite the meagre nectar yields of individual cranberry flowers. 5. Heritable behavioural polymorphisms of the honey bee, such as pollen‐hoarding, can enhance fruit and seed set by a floral host (e.g. cranberry), but only if more preferred pollen hosts are absent or rare. Otherwise, honey bees' broad polylecty, flight range, and daily idiosyncrasies in floral fidelity will obscure specific pollen‐foraging differences at a given floral host, even among paired colonies in a seemingly uniform agricultural setting.     

The use of digital video recorders in pollination biology

1. Digital video recording (DVR) devices, such as the GoPro Hero, have the potential to greatly benefit pollination ecology, but the advantages of digitally recording pollinator activity over direct human observation have not been formally assessed. 2. Two plant taxa, Lavandula angustifolia and Canna ‘sp.’, with differing floral morphology, were used to compare the value of DVR and direct observations in estimating honeybee (Apis mellifera) visitation, flower density and number of flowers visited per foraging bout. 3. The two methods yielded identical results when observing the structurally simple L. angustifolia at both high (10.54 ± 0.52 per plant) and low honeybee density (2.24 ± 0.20 per plant). However, DVR underestimated the number of flowers scored in the field of view (28.7 ± 1.8 direct vs. 22.7 ± 0.9 DVR), the number of honeybees observed (5.3 ± 0.8 direct vs. 3.7 ± 0.7 DVR) and the number of flowers visited during foraging bouts (8.3 ± 1.2 direct vs. 5.5 ± 1.0 DVR) on the more complex Canna ‘sp.’ 4. It is concluded that portable weatherproof DVR devices such as the GoPro Hero are valuable tools for pollination biologists, allowing a single researcher to make simultaneous observations of multiple plants in one or more sites, whilst also allowing the footage to be reviewed. However, DVR devices are limited by their depth and field of view when target plants are large or structurally complex.     

Female and male fitness of a sexually deceptive orchid with a narrow distribution area: from phenotypic traits to spatial distribution patterns

• The Orchidaceae family presents one of the most extravagant pollination mechanisms: deception. While many studies on reproductive success have been performed on food‐deception orchids, less have been performed on sexually deceptive orchids. Here, we focused on Ophrys balearica P. Delforge, an endemic orchid of the Balearic Islands, to study its reproductive ecology, the spatio‐temporal variation of its reproductive success and the individual (floral display and geospatial position) and population parameters (patch size, shape and density) that affect its reproductive success.
• We performed hand‐pollination experiments, along with the recording of floral display parameters and GPS position of over 1,100 individuals from seven populations in two consecutive years. We applied, for the first time, GIS tools to analyse the effects of individual’s position within the population on the reproductive success. Reproductive success was measured both in male (removed pollinia) and female (fruit set) fitness.
• The results confirm that this species is pollinator‐dependent and mostly allogamous, but also self‐compatible. This species showed high values for the cumulative inbreeding depression index and high pollen limitation. Male fitness was almost equal to female fitness between years and populations, and reproductive success exhibited huge spatio‐temporal variation.
• Although we did not find strong correlations between floral display and reproductive success, patches with low‐plant density and individuals in the external portion of the population showed significantly higher plant fitness. These findings must be considered in conservation actions for endangered orchid species, especially considering that most orchids are strongly dependent on pollinators for their species’ fitness.

     

Constructing more informative plant–pollinator networks: visitation and pollen deposition networks in a heathland plant community

Interaction networks are widely used as tools to understand plant–pollinator communities, and to examine potential threats to plant diversity and food security if the ecosystem service provided by pollinating animals declines. However, most networks to date are based on recording visits to flowers, rather than recording clearly defined effective pollination events. Here we provide the first networks that explicitly incorporate measures of pollinator effectiveness (PE) from pollen deposition on stigmas per visit, and pollinator importance (PI) as the product of PE and visit frequency. These more informative networks, here produced for a low diversity heathland habitat, reveal that plant–pollinator interactions are more specialized than shown in most previous studies. At the studied site, the specialization index was lower for the visitation network than the PE network, which was in turn lower than for the PI network. Our study shows that collecting PE data is feasible for community-level studies in low diversity communities and that including information about PE can change the structure of interaction networks. This could have important consequences for our understanding of threats to pollination systems.     

Flowering phenology, fruiting success and progressive deterioration of pollination in an early-flowering geophyte

Spatio-temporal patterns of snowmelt and flowering times affect fruiting success in Erythronium grandiflorum Pursh (Liliaceae) in subalpine western Colorado, USA. From 1990 to 1995, I measured the consistency across years of snowmelt patterns and flowering times along a permanent transect. In most years since 1993, I have monitored fruit set in temporal cohorts (early- to late-flowering groups of plants) at one site. To assess ‘pollination limitation’, I have also conducted supplemental hand-pollination experiments at various times through the blooming season. The onset of blooming is determined by snowmelt, with the earliest years starting a month before the latest years owing to variation in winter snowpack accumulation. Fruit set is diminished or prevented entirely by killing frosts in some years, most frequently but not exclusively for the earlier cohorts. When frosts do not limit fruit set, pollination limitation is frequent, especially in the earlier cohorts. Pollination limitation is strongest for middle cohorts: it tends to be negated by frost in early cohorts and ameliorated by continuing emergence of bumble-bee queens in later cohorts. This lily appears to be poorly synchronized with its pollinators. Across the years of the study, pollination limitation appears to be increasing, perhaps because the synchronization is getting worse.     

SPATIAL FLOWER PARAMETERS AND INSECT SPATIAL VISION

The present article reviews recent and older literature on the spatial parameters that flowers display, as well as on the capacities of anthophilous insects to perceive and use these parameters for optimizing their foraging success. Although co-evolution of plants and pollinators has frequently been discussed with respect to floral colours and insect colour vision, it has rarely been assessed with respect to insect spatial vision and spatial floral cues, such as shape, pattern, size, contrast, symmetry, spatial frequency, contour density and orientation of contours. This review is an attempt to fill this gap. From experimental findings and observations on both flowers and insects, we arrive at the conclusion that all of the spatial and spatio-temporal parameters that flowers offer are relevant to the foraging task and are tuned to the insect's visual capacities and visually guided behaviour. We try, in addition, to indicate that temporal cues are closely related to spatial cues, and must therefore be included when flower–pollinator interactions are examined. We include results that show that colour vision and spatial vision have diverged over the course of evolution, particularly regarding the processing of spatio-temporal information, but that colour vision plays a role in the processing of spatial cues that are independent of temporal parameters. By presenting this review we hope to contribute to closer collaboration among scientists working in the vast fields of botany, ecology, evolution, ethology and sensory physiology.