Bombus terrestris in a mass‐flowering pollinator‐dependent crop: A mutualistic relationship?

Bumblebees (Bombus spp.) rely on an abundant and diverse selection of floral resources to meet their nutritional requirements. In farmed landscapes, mass‐flowering crops can provide an important forage resource for bumblebees, with increased visitation from bumblebees into mass‐flowering crops having an additional benefit to growers who require pollination services. This study explores the mutualistic relationship between Bombus terrestris L. (buff‐tailed bumblebee), a common species in European farmland, and the mass‐flowering crop courgette (Cucurbita pepo L.) to see how effective B. terrestris is at pollinating courgette and in return how courgette may affect B. terrestris colony dynamics. By combining empirical data on nectar and pollen availability with model simulations using the novel bumblebee model Bumble‐BEEHAVE, we were able to quantify and simulate for the first time, the importance of courgette as a mass‐flowering forage resource for bumblebees. Courgette provides vast quantities of nectar to ensure a high visitation rate, which combined with abundant pollen grains, enables B. terrestris to have a high pollination potential. While B. terrestris showed a strong fidelity to courgette flowers for nectar, courgette pollen was not found in any pollen loads from returning foragers. Nonetheless, model simulations showed that early season courgette (nectar) increased the number of hibernating queens, colonies, and adult workers in the modeled landscapes. Synthesis and applications. Courgette has the potential to improve bumblebee population dynamics; however, the lack of evidence of the bees collecting courgette pollen in this study suggests that bees can only benefit from this transient nectar source if alternative floral resources, particularly pollen, are also available to fulfill bees’ nutritional requirements in space and time. Therefore, providing additional forage resources could simultaneously improve pollination services and bumblebee populations.     

Are reward polymorphisms subject to frequency‐ and density‐dependent selection? Evidence from a monoecious species pollinated by deceit

Abstract Most deceit‐pollinated species involve floral dimorphisms characterized by the presence of rewarding male flowers and nonrewarding female flowers. It has been proposed that this polymorphism establishes the conditions for the action of frequency dependent selection (FDS). The tendency of foraging animals to aggregate in areas of high resource density suggests that pollination efficiency and fruit production may be positively influenced by flower density (density dependent selection, DDS). In this paper we offer a graphical model describing the effects of FDS and DDS on a monoecious species pollinated by deceit. We test the FDS and DDS assumptions and the predictions of the model using field observations and experimental populations of Begonia gracilis in which population sex ratio and flower density were controlled. We found a marked effect of both FDS and DDS on pollinator visitation, fruit‐set, and on the probability of female flowers to setting fruits. We conclude that these two types of selection have had a strong influence on the evolution of deceit‐pollinated species.     

Does facilitating pollinator learning impede deceptive orchid attractiveness? A multi‐approach test of avoidance learning

It has often been proposed that nectarless deceptive orchid species exploit naïve pollinators in search of food before they learn to avoid their flowers, and that intraspecific floral trait polymorphism, often noted in this plant group, could prolong the time needed for learning, thus increasing orchid reproductive success. We tested the importance of avoidance learning in a European deceptive orchid, Anacamptis morio, which has been reported to have a highly variable fragrance bouquet among individuals. We used an indirect approach, i.e. we facilitated pollinators’ ability to learn to avoid A. morio by adding anisaldehyde to selected inflorescences, a scent compound that is easily perceived by the natural pollinators and produced in large quantities by the closely related, nectar producing Anacamptis coriophora, a species that shares pollinator species with A. morio. In a series of three experiments (in artificial arrays, in natural populations and in bumblebee behavioural observations), we consistently found no difference either of reproductive success of or visitation rates to scent‐added versus control inflorescences. We also found that the decrease of reproductive success over time in artificial populations of this deceptive species was not as important as expected. Together, these data suggest that pollinators do not fully learn to avoid deceptive inflorescences, and that pollinator avoidance behaviour alone may explain the lower reproductive success usually found in deceptive orchids. We discuss the possible explanations for this pattern in deceptive orchids, particularly in relation to pollinator cognition and learning abilities. Lastly, in light of our results, the potential for higher average reproductive success in deceptive orchids with high phenotypic variability driven by avoidance learning thus appears to be challenged.     

Does the morphological fit between flowers and pollinators affect pollen deposition? An experimental test in a buzz‐pollinated species with anther dimorphism

Some pollination systems, such as buzz‐pollination, are associated with floral morphologies that require a close physical interaction between floral sexual organs and insect visitors. In these systems, a pollinator's size relative to the flower may be an important feature determining whether the visitor touches both male and female sexual organs and thus transfers pollen between plants efficiently. To date, few studies have addressed whether in fact the “fit” between flower and pollinator influences pollen transfer, particularly among buzz‐pollinated species. Here we use Solanum rostratum, a buzz‐pollinated plant with dimorphic anthers and mirror‐image flowers, to investigate whether the morphological fit between the pollinator's body and floral morphology influences pollen deposition. We hypothesized that when the size of the pollinator matches the separation between the sexual organs in a flower, more pollen should be transferred to the stigma than when the visitor is either too small or too big relative to the flower. To test this hypothesis, we exposed flowers of S. rostratum with varying levels of separation between sexual organs, to bumblebees (Bombus terrestris) of different sizes. We recorded the number of visits received, pollen deposition, and fruit and seed production. We found higher pollen deposition when bees were the same size or bigger than the separation between anther and stigma within a flower. We found a similar, but not statistically significant pattern for fruit set. In contrast, seed set was more likely to occur when the size of the flower exceeded the size of the bee, suggesting that other postpollination processes may be important in translating pollen receipt to seed set. Our results suggest that the fit between flower and pollinator significantly influences pollen deposition in this buzz‐pollinated species. We speculate that in buzz‐pollinated species where floral morphology and pollinators interact closely, variation in the visitor's size may determine whether it acts mainly as a pollinator or as a pollen thief (i.e., removing pollen rewards but contributing little to pollen deposition and fertilization).     

Does Neighborhood Floral Display Matter? Fruit Set in Carpenter Bee‐pollinated Heterophragma quadriloculare and Beetle‐pollinated Lasiosiphon eriocephalus

Fruit set is pollen‐limited in the self‐incompatible tree Heterophragma quadriloculare (Bignoniaceae), pollinated by long‐distance flying carpenter bees, and in the self‐compatible shrub Lasiosiphon eriocephalus (Thymdeaceae), pollinated by weak‐flying, sedentary beetles. We studied a single H. quadriloculare population over high and low flowering years and found no difference in bee visitation rates between these years. For H. quadriloculare, neighborhood floral display did not make a significant contribution to reproductive success. We investigated dense and sparse L. eriocephalus populations in the same year. In the low density L. eriocephalus population, individual floral displays were higher than in the dense population, yet reproductive success was low, indicating that plant isolation was a major factor influencing fruit set. This result was due to the extremely low number of beetles per plant and per flower in this population. In the dense L. eriocephalus population, although the displays of individual neighbors were smaller and plants were closer, neighborhood floral display did not contribute significantly to reproductive success, whereas the effect of individual floral display was ambiguous. Species with self‐incompatible rather than self‐compatible breeding systems are expected to experience neighborhood effects on reproductive success; however, at the spatial scale and floral display levels of plants in this study, only individual floral display affected fruit set in H. quadriloculare, whereas neither individual nor neighborhood display influenced fruit set in L. eriocephalus. Therefore, pollinator type, pollinator behavior, and plant and population isolation, rather than breeding system alone, will determine if neighborhood floral display affects fruit set.     

Does egg competition occur in marine broadcast‐spawners?

When the availability of sperm limits female reproductive success, competition for sperm, may be an important broker of sexual selection. This is because sperm limitation can increase the variance in female reproductive success, resulting in strong selection on females to compete for limited fertilization opportunities. Sperm limitation is probably common in broadcast-spawning marine invertebrates, making these excellent candidates for investigating scramble competition between broods of eggs and its consequences for female reproductive success. Here, we report our findings from a series of experiments that investigate egg competition in the sessile, broadcast-spawning polychaete Galeolaria caespitosa. We initially tested whether the order in which eggs encounter sperm affects their fertilization success at two ecologically relevant current regimes. We used a split-clutch-split--ejaculate technique to compare the fertilization success of eggs from individual females that had either first access (competition-free treatment) or second access (egg competition treatment) to a batch of sperm. We found that fertilization success depended on the order in which eggs accessed sperm; eggs that were assigned to the competition-free treatment exhibited significantly higher fertilization rates than those assigned to the egg competition treatment at both current speeds. In subsequent experiments we found that prior exposure of sperm to eggs significantly reduced both the quantity and quality of sperm available to fertilize a second clutch of eggs, resulting in reductions in fertilization success at high and low sperm concentrations. These findings suggest that female traits that increase the likelihood of sperm-egg interactions (e.g. egg size) will respond to selection imposed by egg competition.     

Does intraspecific behavioural variation of pollinator species influence pollination? A quantitative study with hummingbirds and a Neotropical shrub

• Floral visitors differ in their efficacy as pollinators, and the impact of different pollinator species on pollen flow and plant reproduction has been frequently evaluated. In contrast, the impact of intraspecific behavioural changes on their efficacy as pollinators has seldom been quantified.
• We studied a self‐incompatible shrub Palicourea rigida (Rubiaceae) and its hummingbird pollinators, which adjust their behaviour according to floral resource availability. Fluorescence microscopy was used to access pollen tube growth and incompatibility reaction in pistils after a single visit of territorial or intruder hummingbirds in two populations. To characterise the plant populations and possible differences in resource availability between areas we used a three‐term quadrat variance method to detect clusters of floral resources.
• Within‐species variation in foraging behaviour, but not species identity, affected pollinator efficacy. Effectively, hummingbirds intruding into territories deposited more compatible pollen grains on P. rigida stigmas than territory holders in both study areas. Additionally, territory holders deposited more incompatible than compatible pollen grains.
• Our results imply that intraspecific foraging behaviour variation has consequences for pollination success. Quantifying such variation and addressing the implications of intraspecific variability contribute to a better understanding of the dynamics and consequences of plant–pollinator interactions.

     

Does the invasive Lupinus polyphyllus increase pollinator visitation to a native herb through effects on pollinator population sizes?

Invasive plants may compete with native species for abiotic factors as light, space and nutrients, and have also been shown to affect native pollination interactions. Studies have mainly focused on how invasive plants affect pollinator behaviour, i.e. attraction of pollinators to or away from native flowers. However, when an invasive plant provides resources utilized by native pollinators this could increase pollinator population sizes and thereby pollination success in natives. Effects mediated through changes in pollinator population sizes have been largely ignored in previous studies, and the dominance of negative interactions suggested by meta-analyses may therefore be biased. We investigated the impact of the invasive Lupinus polyphyllus on pollination in the native Lotus corniculatus using a study design comparing invaded and uninvaded sites before and after the flowering period of the invasive. We monitored wild bee abundance in transects, and visit rate and seed production of potted Lotus plants. Bumblebee abundance increased 3.9 times in invaded sites during the study period, whereas it was unaltered in uninvaded sites. Total visit rate per Lotus plant increased 2.1 times in invaded sites and decreased 4.4 times in uninvaded sites. No corresponding change in seed production of Lotus was found. The increase in visit rate to Lotus was driven by an increase in solitary bee visitation, whereas mainly bumblebees were observed to visit the invasive Lupinus. The mechanism by which the invasive increases pollinator visit rates to Lotus could be increased availability of other flower resources for solitary bees when bumblebees forage on Lupinus.     

Does a type specimen necessarily or contingently belong to its species?

In a recent article, Alex Levine raises a paradox. It appears that, given some relatively uncontroversial premises about how a species term comes to refer to its species, a type specimen belongs necessarily and contingently to its species. According to Levine, this problem arises if species are individuals rather than natural kinds. I argue that the problem can be generalized: the problem also arises if species are kinds and type specimens are paradigmatic members used to baptize names for species. Indeed, the same problem arises with respect to kinds like gold and the samples used to ground names for them. After arguing that the paradox arises whether or not species are individuals, I attempt to show how the paradox can be resolved. Levine's argument that a type specimen belongs necessarily to its species is specious. The appeal of the argument stems from a failure to distinguish between two different modal statements concerning type specimens, one de dicto and the other de re. Type specimens belong contingently to their respective species. Even so, they can be known a priori to belong to them: hence, that a particular type specimen belongs to its species is an example of contingent a priori knowledge.     

Does ecological filtering across a productivity gradient explain variation in species pool‐richness relationships?

Concerns about the impact of species diversity loss has heightened the importance of determining what limits local diversity and the relative roles of the available species pool and local interactions. Recent theory suggests that local diversity may exhibit linear or asymptotic responses to regional variation in species pools depending resource supply rates. We tested this idea by augmenting species pools in three sites along a natural productivity gradient (a surrogate for resource supply) in a mid-successional old-field. Augmentation of the species pool, enhanced local diversity more at medium than low productivity sites in this field and seed addition had no effect at high productivity, suggesting that diversity was saturated in high resource areas of this field. These results suggest that resource supply rates may mediate species pool-richness relationships. Species additions into cleared plots demonstrated that the observed differences in species recruitment along this gradient were largely driven by abiotic filtering at low and competitive filtering at high productivity. Furthermore, we observed that augmentation of the species pool shifted the productivity–diversity relationship from a negative linear to unimodal suggesting that species pools may influence reported productivity–diversity patterns.     

Does allometry of a sexually selected ornamental trait vary with sexual selection intensity? A multi‐species test in damselflies

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Can we disentangle predator–prey interactions from species distributions at a macro‐scale? A case study with a raptor species

There is a debate on whether an influence of biotic interactions on species distributions can be reflected at macro‐scale levels. Whereas the influence of biotic interactions on spatial arrangements is beginning to be studied at local scales, similar studies at macro‐scale levels are scarce. There is no example disentangling, from other similarities with related species, the influence of predator–prey interactions on species distributions at macro‐scale levels. In this study we aimed to disentangle predator–prey interactions from species distribution data following an experimental approach including a factorial design. As a case of study we selected the short‐toed eagle because of its known specialization on certain prey reptiles. We used presence–absence data at a 100 km2 spatial resolution to extract the explanatory capacity of different environmental predictors (five abiotic and two biotic predictors) on the short‐toed eagle species distribution in peninsular Spain. Abiotic predictors were relevant climatic and topographic variables, and relevant biotic predictors were prey richness and forest density. In addition to the short‐toed eagle, we also obtained the predictor's explanatory capacities for 1) species of the same family Accipitridae (as a reference), 2) for other birds of different families (as controls) and 3) artificial species with randomly selected presences (as null models). We run 650 models to test for similarities of the short‐toed eagle, controls and null models with reference species, assessed by regressions of explanatory capacities. We found higher similarities between the short‐toed eagle and other species of the family Accipitridae than for the other two groups. Once corrected by the family effect, our analyses revealed a signal of predator–prey interaction embedded in species distribution data. This result was corroborated with additional analyses testing for differences in the concordance between the distributions of different bird categories and the distributions of either prey or non‐prey species of the short‐toed eagle. Our analyses were useful to disentangle a signal of predator–prey interactions from species distribution data at a macro‐scale. This study highlights the importance of disentangling specific features from the variation shared with a given taxonomic level.     

Growing with siblings: a common ground for cooperation or for fiercer competition among plants?

Recent work has shown that certain plants can identify their kin in competitive settings through root recognition, and react by decreasing root growth when competing with relatives. Although this may be a necessary step in kin selection, no clear associated improvement in individual or group fitness has been reported to qualify as such. We designed an experiment to address whether genetic relatedness between neighbouring plants affects individual or group fitness in artificial populations. Seeds of Lupinus angustifolius were sown in groups of siblings, groups of different genotypes from the same population and groups of genotypes from different populations. Both plants surrounded by siblings and by genotypes from the same population had lower individual fitness and produced fewer flowers and less vegetative biomass as a group. We conclude that genetic relatedness entails decreased individual and group fitness in L. angustifolius. This, together with earlier work, precludes the generalization that kin recognition may act as a widespread, major microevolutionary mechanism in plants.     

Cancer: A de‐repression of a default survival program common to all cells?

Cancer viewed as a programmed, evolutionarily conserved life-form, rather than just a random series of disease-causing mutations, answers the rarely asked question of what the cancer cell is for, provides meaning for its otherwise mysterious suite of attributes, and encourages a different type of thinking about treatment. The broad but consistent spectrum of traits, well-recognized in all aggressive cancers, group naturally into three categories: taxonomy ("phylogenation"), atavism ("re-primitivization") and robustness ("adaptive resilience"). The parsimonious explanation is not convergent evolution, but the release of an highly conserved survival program, honed by the exigencies of the Pre-Cambrian, to which the cancer cell seems better adapted; and which is recreated within, and at great cost to, its host. Central to this program is the Warburg Effect, whose malign influence permeates well beyond aerobic glycolysis to include biomass interconversion and genomic heuristics. Warburg-type metabolism and genomic instability are targets whose therapeutic disablement is a major priority.     

Population genetics and geometric morphometrics of the <Emphasis Type="Italic">Bombus ephippiatus</Emphasis> species complex with implications for its use as a commercial pollinator

Mexico and Central America are among the most biodiverse regions on Earth, harboring many species with high levels of interpopulation morphological and genetic diversity. The mountainous topography of this region contains isolated sky island habitats that have the potential to promote speciation. This has been studied in vertebrates, yet few studies have examined the phylogeographic and genetic structure of insect species encompassing this region. Here we investigate geographic patterns of genetic and morphological divergence and speciation among widespread populations of the highly polymorphic bumble bee Bombus ephippiatus and its closest relative B. wilmattae. We used DNA sequences from a fragment of cytochrome oxidase I (COI), genotypes for twelve microsatellite markers, and morphometric data from wings to construct a well-supported inference of the divergences among these taxa. We have found complex patterns of genetic isolation and morphological divergence within B. ephippiatus across its geographic range and present evidence that B. ephippiatus comprises multiple independent evolutionary lineages. The pattern of their diversification corresponds to geographic and environmental isolating mechanisms, including the Mexican highlands, the lowlands of the Isthmus of Tehuantepec in southern Mexico, the Nicaraguan Depression, the patchily distributed volcanic ranges in Nuclear Central America and Pleistocene glacial cycles. These results have important implications for the development and distribution of B. ephippiatus as a commercial pollinator in Mexico and Central America.     

Does a multi‐plant diet benefit a polyphagous herbivore? A case study with Bemisia tabaci

Bemisia tabaci (Gennadius) (Hemiptera: Aleyrodidae) is a highly polyphagous herbivore. This research was conducted to compare the development of B. tabaci reared in a multi‐plant treatment (polyculture) with those in single‐plant treatments (monocultures). Adult B. tabaci females fed on a mixture of tomato, cabbage, cotton, cucumber, and kidney bean survived longer and laid more eggs than those fed exclusively on one of these plant species. Egg numbers per plant laid in the polyculture treatment were positively correlated with those laid on the same plant species in the monoculture treatments, and egg numbers per plant laid on tomato, cotton, and cucumber in the polyculture were significantly higher than those laid on the same plants in the monocultures. Concentrations of total protein and trehalose in B. tabaci were not significantly different after 7 days of feeding in the respective treatments, but activities of superoxide dismutases (SOD) and alkaline phosphatase (AKP) of B. tabaci in polyculture were lower than those in monoculture. Conversely, activities of trehalase, sucrase, and amylase in B. tabaci kept in polyculture were higher than those of insects from the monoculture. In each of the monoculture treatments, there was a negative correlation between AKP in B. tabaci and oviposition, and also between AKP and amylase. SOD and sucrase activities in B. tabaci were positively correlated with polyphenol oxidase (PPO) and peroxidase (POD) activities in plants. In the plants damaged by whiteflies in the polyculture treatment, activities of SOD in cucumber, PPO in cotton and kidney bean, and POD in tomato and cucumber were lower than those in the monoculture treatments, whereas SOD in cabbage and catalase (CAT) in tomato in the polyculture treatment were higher than those in the monoculture treatments.     

Does Bt rice pose risks to non‐target arthropods? Results of a meta‐analysis in China

Transgenic Bt rice expressing the insecticidal proteins derived from Bacillus thuringiensis Berliner (Bt) has been developed since 1989. Their ecological risks towards non‐target organisms have been investigated; however, these studies were conducted individually, yielding uncertainty regarding potential agroecological risks associated with large‐scale deployment of Bt rice lines. Here, we developed a meta‐analysis of the existing literature to synthesize current knowledge of the impacts of Bt rice on functional arthropod guilds, including herbivores, predators, parasitoids and detritivores in laboratory and field studies. Laboratory results indicate Bt rice did not influence survival rate and developmental duration of herbivores, although exposure to Bt rice led to reduced egg laying, which correctly predicted their reduced abundance in Bt rice agroecosystems. Similarly, consuming prey exposed to Bt protein did not influence survival, development or fecundity of predators, indicating constant abundances of predators in Bt rice fields. Compared to control agroecosystems, parasitoid populations decreased slightly in Bt rice cropping systems, while detritivores increased. We draw two inferences. One, laboratory studies of Bt rice showing effects on ecological functional groups are mainly either consistent with or more conservative than results of field studies, and two, Bt rice will pose negligible risks to the non‐target functional guilds in future large‐scale Bt rice agroecosystems in China.     

Halobacterium sp. GRB: a species to work with!?

The properties of the halobacterial isolate Halobacterium sp. GRB are discussed, especially in relation to its use as a laboratory strain. Experimental results on this species are described, including the isolation of point mutants in the bacterioopsin gene leading to single amino acid replacements in bacteriorhodopsin, the application of a selection procedure for the isolation of different types of mutants, the genetic stability of Halobacterium sp. GRB and the possibility of isolating a set of isogenic mutants, the conditions for transformation experiments with this species, and specific features of Halobacterium sp. GRB, such as halocin production and the absence of a restriction system, as well as DNA adenosine methylation.     

Does behaviour play a role in house fly resistance to imidacloprid‐containing baits?

The objective of this research was to examine the role and type of behavioural mechanisms that function in house fly, Musca domestica L. (Diptera: Muscidae), resistance to an imidacloprid‐containing commercial fly bait, QuickBayt^®, using an insecticide‐susceptible and an imidacloprid‐resistant strain. Mortality and feeding behaviour were observed through choice bioassays of three post‐imidacloprid selected house fly generations to determine whether flies would consume the bait in the presence of an alternative food source. Mortality rates in choice containers progressively decreased in post‐selection flies as QuickBayt^® no‐choice selections proceeded. There were no differences between the proportions of flies observed contacting QuickBayt^® and sugar, respectively, a finding that eliminates repellency as a mechanism of stimulus‐dependent behavioural resistance. However, differences in QuickBayt^® consumption and subsequent mortality between choice and no‐choice containers provided strong support for the evolution of consumption irritancy‐ or taste aversion‐related behavioural resistance. The results of this study support the responsible rotation of insecticide bait formulations for house fly control.