Pollination Requirements of Almond (Prunus dulcis): Combining Laboratory and Field Experiments.

Almond (Prunus dulcis (Mill.) D. A. Webb; Rosales: Rosaceae) is a cash crop with an estimated global value of over seven billion U.S. dollars annually and commercial varieties are highly dependent on insect pollination. Therefore, the understanding of basic pollination requirements of the main varieties including pollination efficiency of honey bees (Apis mellifera, Linnaeus, Hymenoptera: Apidae) and wild pollinators is essential for almond production. We first conducted two lab experiments to examine the threshold number of pollen grains needed for successful pollination and to determine if varietal identity or diversity promotes fruit set and weight. Further, we examined stigma and ovules of flowers visited by Apis and non-Apis pollinators in the field to study the proportion of almond to non-almond pollen grains deposited, visitation time per flower visit, and tube set. Results indicate that the threshold for successful fertilization is around 60 pollen grains, but pollen can be from any compatible variety as neither pollen varietal identity nor diversity enhanced fruit set or weight. Andrena cerasifolii Cockerell (Hymenoptera: Andrenidae) was a more effective pollinator on a per single visit basis than Apis and syrphid flies. Nevertheless, Apis was more efficient than A. cerasifolii and syrphid flies as they spent less time on a flower during a single visit. Hence, planting with two compatible varieties and managing for both Apis and non-Apis pollinators is likely to be an optimal strategy for farmers to secure high and stable pollination success.

Honey Bee Viruses in Wild Bees: Viral Prevalence, Loads, and Experimental Inoculation.

Evidence of inter-species pathogen transmission from managed to wild bees has sparked concern that emerging diseases could be causing or exacerbating wild bee declines. While some pathogens, like RNA viruses, have been found in pollen and wild bees, the threat these viruses pose to wild bees is largely unknown. Here, we tested 169 bees, representing 4 families and 8 genera, for five common honey bee (Apis mellifera) viruses, finding that more than 80% of wild bees harbored at least one virus. We also quantified virus titers in these bees, providing, for the first time, an assessment of viral load in a broad spectrum of wild bees. Although virus detection was very common, virus levels in the wild bees were minimal-similar to or lower than foraging honey bees and substantially lower than honey bees collected from hives. Furthermore, when we experimentally inoculated adults of two different bee species (Megachile rotundata and Colletes inaequalis) with a mixture of common viruses that is lethal to honey bees, we saw no effect on short term survival. Overall, we found that honey bee RNA viruses can be commonly detected at low levels in many wild bee species, but we found no evidence that these pathogens cause elevated short-term mortality effects. However, more work on these viruses is greatly needed to assess effects on additional bee species and life stages.

Predicting bee community responses to land-use changes: Effects of geographic and taxonomic biases.

Land-use change and intensification threaten bee populations worldwide, imperilling pollination services. Global models are needed to better characterise, project, and mitigate bees' responses to these human impacts. The available data are, however, geographically and taxonomically unrepresentative; most data are from North America and Western Europe, overrepresenting bumblebees and raising concerns that model results may not be generalizable to other regions and taxa. To assess whether the geographic and taxonomic biases of data could undermine effectiveness of models for conservation policy, we have collated from the published literature a global dataset of bee diversity at sites facing land-use change and intensification, and assess whether bee responses to these pressures vary across 11 regions (Western, Northern, Eastern and Southern Europe; North, Central and South America; Australia and New Zealand; South East Asia; Middle and Southern Africa) and between bumblebees and other bees. Our analyses highlight strong regionally-based responses of total abundance, species richness and Simpson's diversity to land use, caused by variation in the sensitivity of species and potentially in the nature of threats. These results suggest that global extrapolation of models based on geographically and taxonomically restricted data may underestimate the true uncertainty, increasing the risk of ecological surprises.

Habitat choice of multiple pollinators in almond trees and its potential effect on pollen movement and productivity: A theoretical approach using the Shigesada-Kawasaki-Teramoto model.

California's almond industry, valued at $2.3 billion per year, depends on the pollinator services of honey bees, although pollination by other insects, mainly solitary wild bees, is being investigated as an alternative because of recent declines in the number of honey bee colonies. Our objective is to model the movements of honey bees and determine the conditions under which they will forage in less favorable areas of a tree and its surroundings when other pollinators are present. We hypothesize that foraging in less favorable areas leads to increased movement between trees and increased cross pollination between varieties which is required for successful nut production. We use the Shigesada-Kawasaki-Teramoto model (1979) which describes the density of two species in a two-dimensional environment of variable favorableness with respect to intrinsic diffusions and intra and interspecific interactions of species. The model is applied to almond pollination by honey bees and other pollinators with environmental favorableness based on the distribution of flowers in trees. Using the spectral-Galerkin method in a rectangular domain, we numerically approximated the two-dimensional nonlinear parabolic partial differential system arising in the model. When cross-diffusion or interspecific effects of other pollinators was high, honey bees foraged in less favorable areas of the tree. In the model, high cross-diffusion also resulted in increased activity in honey bees which manifested itself in the field in terms of accelerations, decelerations, and changes in direction, indicating rapid redistribution of densities to an equilibrium state. Empirical analysis of the number of honey bees and other visitors in 2-min intervals to almond trees shows a negative relationship, indicating cross-diffusion effects in nature with the potential to increase movement to a different tree with a more favorable environment, potentially increasing nut production.

Reproduction of Amorpha canescens (Fabaceae) and diversity of its bee community in a fragmented landscape.

Loss of insect pollinators due to habitat fragmentation often results in negative effects on plant reproduction, but few studies have simultaneously examined variation in the bee community, site characteristics and plant community characteristics to evaluate their relative effects on plant reproduction in a fragmented habitat. We examined the reproduction of a common tallgrass prairie forb, Amorpha canescens (Fabaceae), in large (>40 ha) and small (<2 ha) prairie remnants in Iowa and Minnesota in relation to the diversity and abundance of its bee visitors, plant population size, and species density of the forb flowering community. We found significant positive effects of the diversity of bees visiting A. canescens on percent fruit set at a site in both years of the study and in 2002 an additional significant positive effect of plant species density. Abundance of bees visiting A. canescens had a significant positive effect on percent fruit set in 2002, but was only marginally significant in 2003. In 2003 but not 2002, the plant species density at the sites had a significant negative effect on the diversity and abundance of bees visiting A. canescens, indicating community-level characteristics can influence the bee community visiting any one species. Site size, a common predictor of plant reproduction in fragmented habitats did not contribute to any models of fruit set and was only marginally related to bee diversity one year. Andrena quintilis, one of the three oligolectic bee species associated with A. canescens, was abundant at all sites, suggesting it has not been significantly affected by fragmentation. Our results show that the diversity of bees visiting A. canescens is important for maintaining fruit set and that bee visitation is still sufficient for at least some fruit set in all populations, suggesting these small remnants act as floral resource oases for bees in landscapes often dominated by agriculture.

Are seedlings from small seeds always inferior to seedlings from large seeds? Effects of seed biomass on seedling growth in Pastinaca sativa L.

Previous field studies of recruitment in Pastinaca sativa L. indicate that more seedlings from small seeds than from large seeds survive short-term droughts. To explore this phenomenon, the effects of variation in seed biomass in Pastinaca sativa on embryo size and seedling characteristics 10, 20, 30 and 40 days after emergence were investigated. On the basis of most characteristics, seedlings from large seeds should be superior to seedlings from small seeds. Embryo length and cotyledon area were positively related to seed biomass, as were above ground biomass, total leaf area, and root biomass in all harvests. Total seedling biomass was positively related to seed biomass in the 10, 20, and 30 day harvests, but not the 40 day harvest. However, the ratio of maximum root length/total leaf area was negatively related to seed biomass in the 10 and 20 day harvests, suggesting that, under drought conditions, seedlings from small seeds may transpire less water than those from large seeds relative to their ability to reach water supplies. Although seedlings from larger seeds had greater root biomass, this may be of little advantage under drought conditions since approximately 90% of the root biomass is in the upper 10 cm of soil which dries out quickly. The advantage that seedlings from small seeds have under drought conditions is short-lived, lasting about 20 days in the glasshouse and an estimated 60-90 days in the field. This advantage is short-term because the relationship between seed biomass and resource allocation patterns changes during early seedling development.

Consequences of a mixed reproductive system in the hog peanut, Amphicarpaea bracteata, (Fabaceae).

The forest annual, Amphicarpaea bracteata L. can reproduce via aerial chasmogamous, aerial cleistogamous, and subterranean cleistogamous flowers. Both plant size and light intensity influenced the utilization of the three modes of reproduction. chasmogamous and aerial cleistogamous flower number and the ratio of chasmogamous flowers to the total number of aerial flowers increased with plant size. The latter demonstrated a shift to xenogamy and outbreeding in larger plants. Light intensity indirectly influenced reproductive modes through its infuence on plant size. Seed set by both types of aerial flowers was low and unrelated to plant size. Subterranean seed number and the total dry weight of subterranean seeds per plant increased with size. The subterranean seeds of Amphicarpaea bracteata are thirty-four times larger than the aerial seeds (fresh weight). Under field conditions, subterranean seeds had greater germination after one year than acrial seeds. The plants arising from subterranean seeds were significantly larger and more fecund than those from aerial seeds. Seeds produced by aerial cleistogamous, hand selfpollinated chasmogamous, and naturally pollinated chasmogamous flowers had equivalent germination rates and produced plants of equal size and fecundity. This suggests that the outbred progeny from chasmogamous flowers have no advantage over the inbred progeny from aerial cleistogamous flowers.

Plant-herbivore interactions: Insect induced changes in host plant sex expression and fecundity.

We investigated changes in the reproduction of Pastinaea sativa which result in compensatory reproduction when herbivore feeding destroys the first infloresence produced. Late developing infloresences of damaged plants produce more flowers and a greater proportion of hermaphroditic flowers than late developing infloresences of control plants. In damaged plants the proportion of hermaphroditic flowers producing seeds in late developing infloresences also increases. These effects are due to the mechanical damage resulting from larval feeding. The herbivore acts as a mediator in a number of important plant processes affecting fecundity, including expression of sex.

Compensatory reproduction in a biennial herb following insect defloration.

The ability of the biennial herb, Pastinaca sativa L. (wild parsnip), to respond to and compensate for destruction of primary umbel seeds by the larvae of Depressaria pastinacella (Lepidoptera: Oecophoridae) was analyzed by comparing umbel and seed production of damaged and undamaged plants collected from five populations. Plants with a basal stem diameter smaller than 8 mm suffer a reduction in seed set of about 50% when the primary umbel is destroyed but larger plants are able to compensate for loss of primary umbel seeds by increased seed set of tertiary umbels. Depending on plant size, this is due to either an increase in the number of tertiary umbels that reach maturity or an increase in the number of seeds per tertiary umbel. Although seeds of tertiary umbels are significantly smaller than those of primary or secondary umbels, their viability is equivalent to that of secondary seeds and may be greater than that of primary seeds. Characteristics of P. sativa's reproduction, such as the long flowering period and the initiation of more umbels than the plant is normally able to bring to maturity, are important to P. sativa's ability to compensate for the effects of herbivore damage.

The resistance of Pteridium aquilinum (L.) Kuhn to insect attack by Trichoplusia ni (Hübn.).

Resistance of Pteridium aquilinum to insect attack was studied by incorporating air dried bracken leaf meal and extracts of bracken leaf meal into an artificial diet for Trichoplusia ni (Lepidoptera: Noctuidae). When larvae are reared on diets containing 6% bracken leaf meal, they do not mature past the second instar and after 16 days the average weight is approximately 1 mg compared to 70 mg for control larvae. Feeding initiation studies indicate that a feeding deterrent is present in bracken fern but feeding rates and food utilization efficiency studies suggest that either the deterrent or another compound also functions as a toxin. This toxin does not affect growth, feeding rates, or utilization efficiency for the first 4 days after third instar larvae are transferred to a diet containing the water extract of bracken leaf meal; thereafter growth is terminated and feeding is greatly reduced. The active factor is water soluble, heat labile, and non-volatile and this partial characterization indicates that neither the bracken ecdysones or the anti-thiamine factor of bracken is involved in the resistance of this fern to insect attack by T. ni.