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What is pollination and what are pollinators in agriculture?
Abstract
This chapter provides an overview of the history of pollination biology, it begins by discussing the basics of pollination and goes on to discuss pollinators and their diversity. Sections also cover the ecology and evolution of floral traits, domestication and its impact on plant-pollinator relationships and how pollinators can impact agriculture. A section on modern agriculture and pollinators is also provided.
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Pollinator decline has attracted global attention and substantial efforts are underway to respond through national pollinator strategies and action plans. These policy responses require clarity on what is driving pollinator decline and what risks it generates for society in different parts of the world. Using a formal expert elicitation process, we evaluated the relative regional and global importance of eight drivers of pollinator decline and ten consequent risks to human well-being. Our results indicate that global policy responses should focus on reducing pressure from changes in land cover and configuration, land management and pesticides, as these were considered very important drivers in most regions. We quantify how the importance of drivers and risks from pollinator decline, differ among regions. For example, losing access to managed pollinators was considered a serious risk only for people in North America, whereas yield instability in pollinator-dependent crops was classed as a serious or high risk in four regions but only a moderate risk in Europe and North America. Overall, perceived risks were substantially higher in the Global South. Despite extensive research on pollinator decline, our analysis reveals considerable scientific uncertainty about what this means for human society.
Deformed wing virus (DWV) is a persistent pathogen of European honey bees and the major contributor to overwintering colony losses. The prevalence of DWV in honey bees has led to significant concerns about spillover of the virus to other pollinating species. Bumble bees are both a major group of wild and commercially-reared pollinators. Several studies have reported pathogen spillover of DWV from honey bees to bumble bees, but evidence of a sustained viral infection characterized by virus replication and accumulation has yet to be demonstrated. Here we investigate the infectivity and transmission of DWV in bumble bees using the buff-tailed bumble bee Bombus terrestris as a model. We apply a reverse genetics approach combined with controlled laboratory conditions to detect and monitor DWV infection. A novel reverse genetics system for three representative DWV variants, including the two master variants of DWV-type A and B-was used. Our results directly confirm DWV replication in bumble bees but also demonstrate striking resistance to infection by certain transmission routes. Bumble bees may support DWV replication but it is not clear how infection could occur under natural environmental conditions.
Most of the world’s crops depend on pollinators, so declines in both managedand wild bees raise concerns about food security. However, the degree towhich insect pollination is actually limiting current crop production ispoorly understood, as is the role of wild species (as opposed to managed hon-eybees) in pollinating crops, particularly in intensive production areas. Weestablished a nationwide study to assess the extent of pollinator limitation inseven crops at 131 locations situated across major crop-producing areas ofthe USA. We found that five out of seven crops showed evidence of pollinatorlimitation. Wild bees and honeybees provided comparable amounts of pollina-tion for most crops, even in agriculturally intensive regions. We estimated thenationwide annual production value of wild pollinators to the seven crops westudied at over $1.5 billion; the value of wild bee pollination of all pollinator-dependent crops would be much greater. Our findings show that pollinatordeclines could translate directly into decreased yields or production formost of the crops studied, and that wild species contribute substantially topollination of most study crops in major crop-producing regions.
Human populations and their use of land have reshaped landscapes for thousands of years, creating the anthropogenic biomes (anthromes) that now cover most of the terrestrial biosphere. Here we introduce the first global reconstruction and mapping of anthromes and their changes across the 12,000-year interval from 10,000 BCE to 2015 CE; the Anthromes 12K dataset. Anthromes were mapped using gridded global estimates of human population density and land use from the History of the Global Environment database (HYDE version 3.2) by a classification procedure similar to that used for prior anthrome maps. Anthromes 12K maps generally agreed with prior anthrome maps for the same time periods, though significant differences were observed, including a substantial reduction in Rangelands anthromes in 2000 CE but with increases before that time. Differences between maps resulted largely from improvements in HYDE’s representation of land use, including pastures and rangelands, compared with the HYDE 3.1 input data used in prior anthromes maps. The larger extent of early land use in Anthromes 12K also agrees more closely with empirical assessments than prior anthrome maps; the result of an evidence-based paradigm shift in characterizing the history of Earth’s transformation through land use, from a mostly recent large-scale conversion of uninhabited wildlands, to a long-term trend of increasingly intensive transformation and use of already inhabited and used landscapes. The spatial history of anthropogenic changes depicted in Anthromes 12K remain to be validated, especially for earlier time periods. Nevertheless, Anthromes 12K is a major advance over all prior anthrome datasets and provides a new platform for assessing the long-term environmental consequences of human transformation of the terrestrial biosphere.
Pollinator nutritional ecology provides insights into plant–pollinator interactions, coevolution, and the restoration of declining pollinator populations. Bees obtain their protein and lipid nutrient intake from pollen, which is essential for larval growth and development as well as adult health and reproduction. Our previous research revealed that pollen protein to lipid ratios (P:L) shape bumble bee foraging preferences among pollen host-plant species, and these preferred ratios link to bumble bee colony health and fitness. Yet, we are still in the early stages of integrating data on P:L ratios across plant and bee species. Here, using a standard laboratory protocol, we present over 80 plant species’ protein and lipid concentrations and P:L values, and we evaluate the P:L ratios of pollen collected by three bee species. We discuss the general phylogenetic, phenotypic, behavioral, and ecological trends observed in these P:L ratios that may drive plant–pollinator interactions; we also present future research questions to further strengthen the field of pollination nutritional ecology. This dataset provides a foundation for researchers studying the nutritional drivers of plant–pollinator interactions as well as for stakeholders developing planting schemes to best support pollinators.
Despite the paramount role of floral fragrance in pollinator attraction and reproduction in flowering plants, we know little about its evolution under natural conditions. Here we show that by reducing herbivore load with pesticide application, plants showed evolutionary changes in their floral fragrance within 4 generations when compared to plants with natural levels of herbivory. We interpret this finding in the context of relaxed physiological and ecological trade-offs with fewer herbivores, potentially facilitating selection by bees on specific aromatic volatiles. Our study confirms earlier findings of experimental evolution under highly controlled and simplified conditions in the greenhouse.
Native bees are in decline as many species are sensitive to habitat loss, climate change, and non-target exposure to synthetic pesticides. Recent laboratory and semi-field assessments of pesticide impacts on bees have focused on neonicotinoid insecticides. However, field studies evaluating influences of neonicotinoid seed treatments on native bee communities of North America are absent from the literature. On four Conservation Areas of Missouri, we sampled row-cropped (treated, n = 15) and reference (untreated, n = 9) agricultural fields, and their surrounding field margins for neonicotinoids in soil and non-target vegetation (i.e., native wildflowers). Wildflowers were further collected and screened for the presence of fungicides. Concurrently, we sampled native bees over three discrete time points throughout the agricultural growing season to assess potential impacts of seed treatment use on local bee populations over time. Neonicotinoids were detected in 87% to 100% of treated field soils and 22% to 56% of reference field soils. In adjacent field margin soils, quantifiable concentrations were measured near treated (53% to 93% detection) and untreated fields (33% to 56% detection). Fungicides were detected in < 40% of wildflowers, whereas neonicotinoids were rarely detected in field margin vegetation (< 7%). Neonicotinoid concentrations in margin soils were negatively associated with native bee richness (β = −0.21, P < 0.05). Field margins with a combination of greater neonicotinoid concentrations in soil and fungicides in wildflowers also contained fewer wild bee species (β = −0.21, P < 0.001). By comparison, bee abundance was positively influenced by the number of wildflower species in bloom with no apparent impact of pesticides. Results of this study indicate that neonicotinoids in soil are a potential route of exposure for pollinator communities, specifically ground-nesting species. Importantly, native bee richness in non-target field margins may be negatively affected by the use of neonicotinoid seed treatments in agroecosystems.
Pathogen spillover from managed bees is increasingly considered as a possible cause of pollinator decline. Though spillover has been frequently documented, evidence of the pathogen's virulence in the new host or mechanism of transmission is rare. Stingless bees (Apocrita: Meliponini) are crucial pollinators pan-tropically and overlap with managed honeybees (Apis mellifera) in much of their range. Nosema ceranae is the most prevalent disease of adult A. mellifera. We used laboratory experiments and field surveys to investigate the susceptibility of stingless bees (Tetragonula hockingsi) to N. ceranae, infection prevalence and transmissibility via flowers. We found that 67% of T. hockingsi fed sucrose with N. ceranae had detectable spores in their ventriculus, and they died at 2.96 times the rate of sucrose-only fed bees. Five of six field hives harboured bees with N. ceranae present at least once during our five-month survey, with prevalence up to 20%. In our floral transmission experiment, 67% of inflorescences exposed to infected A. mellifera yielded N. ceranae spores, and all resulted in T. hockingsi with N. ceranae spores in their guts. We conclude that N. ceranae is virulent in T. hockingsi under laboratory conditions, is common in the local T. hockingsi population and is transmissible via flowers.
Non-native plant species reliant on insect pollination must attract novel pollinators in their introduced habitat to reproduce. Indeed, pollination services provided by resident floral visitors may contribute to the spread of non-native species, which may then affect the pollination services received by native plants. To determine the mechanisms by which an invasive thistle attracts pollinators in its introduced range, and whether its presence changes the pollinator visitation to native plant species, we compared bee visitation to native plants in the presence or absence of the invader. We experimentally tested the effect of a thistle invasion into a native plant community. We found that the non-native thistle was the most attractive of the plant species to visiting bee species. However, there was no effect of experimental treatment (presence of thistle) on bee abundance or visitation rate (bees per unit floral area per sample) to native plant species. Across 68 bee and 6 plant species, we found a significant correlation between pollen protein content and bee abundance and visitation rate. Thistle pollen also had a similar protein:lipid ratio to legumes, which correlated with bumble bee visitation. The high protein content of the thistle pollen, as compared to four native asters, may allow it to attract pollinators in novel ecosystems, and potentially contribute to its success as an invader. At the same time, this high protein pollen may act as a novel resource to pollinators in the thistle’s invaded range.
Beetles (Coleoptera) are a diverse group of overlooked pollinators, considered particularly important in tropical ecosystems. The role of the most diverse beetle family, Staphylinidae, as pollinators is generally considered minor, yet their relationships
with plants are mostly unknown. Although often referred to as opportunistic visitors, it is arguable that the true extent of rove beetle pollination is underestimated given their frequency of visitation to flowers. This review comprehensively analysed the
plant–pollinator or visitor interactions of the Staphylinidae and uncovered 108 well-described staphylinid–flower interactions across 27 seed plant families. Of these interactions, Staphylinidae were considered either potential or conclusive pollinators
for 56 plant species, having either a primary or secondary role in pollination. Conversely, Staphylinidae were visitors to 40 plant species with a negligible role in pollination. For the remaining 12 interactions and additional anecdotal reports, the role of staphylinids as pollinators was unresolved. Staphylinid–flower interactions were most prevalent in the monocots and magnoliids (families: Araceae, Annonaceae, Arecaceae, and Magnoliaceae) involving predominantly generalist pollination systems, and interactions were limited to six staphylinid subfamilies (Omaliinae, Tachyporinae, Aleocharinae, Oxytelinae,
Paederinae, and Staphylininae). Trends in the involvement of staphylinid subfamilies with particular plant lineages were identified, associated with differences in insect habit and floral rewards. Overall this review indicates that the role of Staphylinidae as pollinators, and Coleoptera as a whole, is underestimated. Caution, however, must be given to inferring the role of staphylinids in pollination because rove beetles commonly function as inadvertent secondary pollinators or antagonists there to fulfil other ecological roles.
Bombus impatiens, the common eastern bumble bee, is the first bumble bee established outside of its native range in North America. Native to the eastern portion of the continent, the species was imported to British Columbia in the early 2000s for greenhouse pollination and subsequently became established in the wild. Here we report on the continuing expansion of its range in the Pacific Northwest, including the detection of gynes and workers in Washington State. Sightings of B. impatiens in the region have become increasingly common based on various Internet mapping and reporting sites. The species has been observed about 30 km south of the British Columbia border, or 60 km from the first British Columbia detections. Species distribution models indicate that the Puget Sound and Willamette Valley are suitable habitat, and the bee will likely continue to expand its range southward towards California. The potential impacts of B. impatiens in the region are unknown and will be monitored in future research.
Pollination is a critical ecosystem service underpinning the productivity of agricultural systems across the world. Wild insect populations provide a substantial contribution to the productivity of many crops and seed set of wild flowers. However, large-scale evidence on species-specific trends among wild pollinators are lacking. Here we show substantial inter-specific variation in pollinator trends, based on occupancy models for 353 wild bee and hoverfly species in Great Britain between 1980 and 2013. Furthermore, we estimate a net loss of over 2.7 million occupied 1 km² grid cells across all species. Declines in pollinator evenness suggest that losses were concentrated in rare species. In addition, losses linked to specific habitats were identified, with a 55% decline among species associated with uplands. This contrasts with dominant crop pollinators, which increased by 12%, potentially in response agri-environment measures. The general declines highlight a fundamental deterioration in both wider biodiversity and non-crop pollination services.
There is growing concern that some bee populations are in decline, potentially threatening pollination security in agricultural and non-agricultural landscapes. Among the numerous causes associated with this trend, nutritional stress resulting from a mismatch between bee nutritional needs and plant community provisioning has been suggested as one potential driver. To ease nutritional stress on bee populations in agricultural habitats, agri-environmental protection schemes aim to provide alternative nutritional resources for bee populations during times of need. However, such efforts have focused mainly on quantity (providing flowering plants) and timing (during flower-scarce periods), while largely ignoring the quality of the offered flower resources. In a first step to start addressing this information gap, we have used literature data to compile a comprehensive geographically explicit dataset on nectar quality (i.e., total sugar concentration), offered to bees both within fields (crop and weed species) as well as outside fields (wild species) around the globe. Social bees are particularly sensitive to nectar sugar concentrations, which directly impact calorie influx into the colony and consequently their fitness making it an important resource quality marker. We find that the total nectar sugar concentrations in general do not differ between the three plant communities studied. In contrast we find increased variability in nectar quality in the wild plant community compared to crop and weed community, which is likely explained by the increased phylogenetic diversity in this category of plants. In a second step we explore the influence of local habitat on nectar quality and its variability utilizing a detailed sunflower (Helianthus annuus L.) data set and find that geography has a small, but significant influence on these parameters. In a third step we identify crop groups (genera), which provide sub-optimal nectar resources for bees and suggest high quality alternatives as potential nectar supplements. In the long term this data set could serve as a starting point to systematically collect more quality characteristics of plant provided resources to bees, which ultimately can be utilized by scientist, regulators, NGOs and farmers to improve the flower resources offered to bees. We hope that ultimately this data will help to ease nutritional stress for bee populations and foster a data informed discussion about pollinator conservation in modern agricultural landscapes.
The timing of the origin of angiosperms is a hotly debated topic in plant evolution. Molecular dating analyses that consistently retrieve pre‐Cretaceous ages for crown‐group angiosperms have eroded confidence in the fossil record, which indicates a radiation and possibly also origin in the Early Cretaceous. Here, we evaluate paleobotanical evidence on the age of the angiosperms, showing how fossils provide crucial data for clarifying the situation. Pollen floras document a Northern Gondwanan appearance of monosulcate angiosperms in the Valanginian and subsequent poleward spread of monosulcates and tricolpate eudicots, accelerating in the Albian. The sequence of pollen types agrees with molecular phylogenetic inferences on the course of pollen evolution, but it conflicts strongly with Triassic and early Jurassic molecular ages, and the discrepancy is difficult to explain by geographic or taphonomic biases. Critical scrutiny shows that supposed pre‐Cretaceous angiosperms either represent other plant groups or lack features that might confidently assign them to the angiosperms. However, the record may allow the Late Jurassic existence of ecologically restricted angiosperms, like those seen in the basal ANITA grade. Finally, we examine recently recognized biases in molecular dating and argue that a thoughtful integration of fossil and molecular evidence could help resolve these conflicts.
Floral chemistry mediates plant interactions with herbivores, pathogens, and pollinators. The chemistry of floral nectar and pollen, the primary food rewards for pollinators, can affect both plant reproduction and pollinator health. Although the existence and functional significance of nectar and pollen secondary metabolites has long been known, comprehensive quantitative characterizations of secondary chemistry exist for only a few species. Moreover, little is known about intraspecific variation in nectar and pollen chemical profiles. Because the ecological effects of secondary chemicals are dose‐dependent, heterogeneity across genotypes and populations could influence floral trait evolution and pollinator foraging ecology. To better understand within‐ and across‐species heterogeneity in nectar and pollen secondary chemistry, we undertook exhaustive LC‐MS and LC‐UV‐based chemical characterizations of nectar and pollen methanol extracts from 31 cultivated and wild plant species. Nectar and pollen were collected from farms and natural areas in Massachusetts, Vermont, and California, USA, in 2013 and 2014. For wild species, we aimed to collect 10 samples from each of three sites. For agricultural and horticultural species, we aimed for 10 samples from each of three cultivars. Our data set (1,535 samples, 102 identified compounds) identifies and quantifies each compound recorded in methanolic extracts, and includes chemical metadata that describe the molecular mass, retention time, and chemical classification of each compound. A reference phylogeny is included for comparative analyses. We found that each species possessed a distinct chemical profile; moreover, within species, few compounds were found in both nectar and pollen. The most common secondary chemical classes were flavonoids, terpenoids, alkaloids and amines, and chlorogenic acids. The most common compounds were quercetin and kaempferol glycosides. Pollens contained high concentrations of hydroxycinnamoyl‐spermidine conjugates, mainly triscoumaroyl and trisferuloyl spermidine, found in 71% of species. When present, pollen alkaloids and spermidines had median nonzero concentrations of 23,000 μmol/L (median 52% of recorded micromolar composition). Although secondary chemistry was qualitatively consistent within each species and sample type, we found significant quantitative heterogeneity across cultivars and sites. These data provide a standard reference for future ecological and evolutionary research on nectar and pollen secondary chemistry, including its role in pollinator health and plant reproduction. Data are published under a Creative Commons Attribution License (CC BY 3.0 US) and may be freely used if properly cited.
Loss of seed shattering is a key trait in crop domestication, particularly for grain crops. For wild plants, seed shattering is a crucial mechanism to achieve greater fitness, although in the agricultural context, this mechanism reduces harvesting efficiency, especially under dry conditions. Loss of seed shattering was acquired independently in different monocotyledon and dicotyledon crop species by ‘convergent phenotypic evolution’, leading to similar low dehiscent and indehiscent phenotypes. Here, the main aim is to review the current knowledge about seed shattering in crops, in order to highlight the tissue modifications that underlie the convergent phenotypic evolution of reduced shattering in different types of fruit, from the silique of Brassicaceae species, to the pods of legumes and spikes of cereals. Emphasis is given to legumes, with consideration of recent data obtained for the common bean. The current review also discusses to what extent convergent phenotypes arose from parallel changes at the histological and/or molecular levels. For this reason, an overview is included of the main findings relating to the genetic control of seed shattering in the model species Arabidopsis thaliana and in other important crops
The high pollen and nectar yield of Asteraceae flowers combined with the abundance and diversity of these plants should make them an important pollen source for both specialist and generalist bees. However, studies have found Asteraceae pollen to be a poor diet for some generalist bees, possibly owing to nutrient deficiencies or the presence of secondary metabolites or digestive barriers that prevent nutrient assimilation. Mixing pollens could allow bees to exploit these unfavourable pollens, if the different pollen sources help to complement nutrient deficiencies or alleviate the effects of toxic secondary metabolites. In our study, we examined how the proportion of sunflower (Helianthus annuus, Asteraceae) pollen in the diet of captive-reared bumblebees (Bombus impatiens) affects the survival of bees in queenless microcolonies. Bees fed on sunflower pollen had significantly shorter lifespans than bees fed on broad bean (Vicia faba, Fabaceae), rapeseed (Brassica napus, Brassicaceae) or Cucurbitaceae pollen. However, survival on mixed pollen diets containing 50% sunflower pollen was as great as that on non-sunflower diets, which suggests that the other pollens were able to compensate for the low nutritive quality of the sunflower pollen. Due to agricultural intensification and a loss of wildflowers, farmland monocultures (e.g. sunflower crops) can be important floral resources for bumblebees. Our study suggests that providing alternative floral resources of high nutritive quality could help mitigate the potential harmful effects of a monofloral sunflower diet on bumblebees.
The benefits of insect pollination to crop yields depend on genetic and environmental factors including plant self-fertility, pollinator visitation rates, and pollinator efficacy. While many crops benefit from insect pollination, such variation in pollinator benefits across both plant cultivars and growing regions is not well documented. In this study, across three states in the northern Great Plains, United States, from 2016 to 2017, we evaluated the pollinator-mediated yield increases for 10 varieties of confection sunflowers, Helianthus annuus L. (Asterales: Asteraceae), a plant that is naturally pollinator-dependent but was bred for self-fertility. We additionally measured pollinator visitation rates and compared per-visit seed set across pollinator taxa in order to determine the most efficacious sunflower pollinators. Across all locations and hybrids, insect pollination increased sunflower yields by 45%, which is a regional economic value of over 56 million. There was, however, some variation in the extent of pollinator benefits across locations and plant genotypes, and such variation was significantly related to pollinator visitation rates, further highlighting the value of pollinators for confection sunflowers. Female Andrena helianthi Robertson (Hymenoptera: Andrenidae) and Melissodes spp. (Hymenoptera: Apidae) were the most common and effective pollinators, while other bees including managed honey bees (Hymenoptera: Halictidae), Apis mellifera L. (Hymenoptera: Apidae), small-bodied sweat bees (Hymenoptera: Halictidae), bumble bees Bombus spp. (Hymenoptera: Apidae), and male bees were either infrequent or less effective on a per-visit basis. Our results illustrate that wild bees, in particular the sunflower specialists A. helianthi and Melissodes spp., provide significant economic benefits to confection sunflower production.
Crop domestication can lead to weakened expression of plant defences, with repercussions for herbivore and pathogen susceptibility. However, little is known about how domestication alters traits that mediate other important ecological interactions in crops, such as pollination. Secondary metabolites, which underpin many defence responses in plants, also occur widely in nectar and pollen and influence plant-pollinator interactions. Thus, domestication may also affect secondary compounds in floral rewards, with potential consequences for pollinators. To test this hypothesis, we chemically analysed nectar and pollen from wild and cultivated plants of highbush blueberry (Vaccinium corymbosum L.), before conducting an artificial diet bioassay to examine pollinator-pathogen interactions. Our results indicated that domestication has significantly altered the chemical composition of V. corymbosum nectar and pollen, and reduced pollen chemical diversity in cultivated plants. Of 20 plant metabolites identified in floral rewards, 13 differed significantly between wild and cultivated plants, with a majority showing positive associations with wild compared to cultivated plants. These included the amino acid phenylalanine (4.5 times higher in wild nectar, 11 times higher in wild pollen), a known bee phagostimulant and essential nutrient; and the antimicrobial caffeic acid ester 4-O-caffeoylshikimic acid (two times higher in wild nectar). We assessed the possible biological relevance of variation in caffeic acid esters in bioassays, using the commercially available 3-O-caffeoylquinic acid. This compound reduced Bombus impatiens infection by a prominent gut pathogen (Crithidia) at concentrations that occurred in wild but not cultivated plants, suggesting that domestication may influence floral traits with consequences for bee health. Appreciable levels of genetic variation and heritability were found for most floral reward chemical traits, indicating good potential for selective breeding. Our study provides the first assessment of plant domestication effects on floral reward chemistry and its potential repercussions for pollinator health. Given the central importance of pollinators for agriculture, we discuss the need to extend such investigations to pollinator-dependent crops more generally and elaborate on future research directions to ascertain wider trends, consequences for pollinators, mechanisms, and breeding solutions.
Global declines in pollinators, including bees, can have major consequences for ecosystem services. Bees are dominant pollinators, making it imperative to mitigate declines. Pathogens are strongly implicated in the decline of native and honey bees. Diet affects bee immune responses, suggesting the potential for floral resources to provide natural resistance to pathogens. We discovered that sunflower (Helianthus annuus) pollen dramatically and consistently reduced a protozoan pathogen (Crithidia bombi) infection in bumble bees (Bombus impatiens) and also reduced a microsporidian pathogen (Nosema ceranae) of the European honey bee (Apis mellifera), indicating the potential for broad anti-parasitic effects. In a field survey, bumble bees from farms with more sunflower area had lower Crithidia infection rates. Given consistent effects of sunflower in reducing pathogens, planting sunflower in agroecosystems and native habitat may provide a simple solution to reduce disease and improve the health of economically and ecologically important pollinators.
The domestication syndrome of many plants includes changes in their mating systems. The evolution of the latter is shaped by ecological and genetic factors that are particular to an area. Thus, the reproductive biology of wild relatives must be studied in their natural distribution to understand the mating system of a crop species as a whole. Gossypium hirsutum (upland cotton) includes both domesticated varieties and wild populations of the same species. Most studies on mating systems describe cultivated cotton as self-pollinated, while studies on pollen dispersal report outcrossing; however, the mating system of upland cotton has not been described as mixed and little is known about its wild relatives. In this study we selected two wild metapopulations for comparison with domesticated plants and one metapopulation with evidence of recent gene flow between wild relatives and the crop to evaluate the mating system of cotton’s wild-to-domesticated complex. Using classic reproductive biology methods, our data demonstrate that upland cotton presents a mixed mating system throughout the complex. Given cotton’s capacity for outcrossing, differences caused by the domestication process in cultivated individuals can have consequences for its wild relatives. This characterization of the diversity of the wild relatives in their natural distribution, as well as their interactions with the crop, will be useful to design and implement adequate strategies for conservation and biosecurity.
Managed bees are critical for crop pollination worldwide. As the demand for pollinator-dependent crops increases, so does the use of managed bees. Concern has arisen that managed bees may have unintended negative impacts on native wild bees, which are important pollinators in both agricultural and natural ecosystems. The goal of this study was to synthesize the literature documenting the effects of managed honey bees and bumble bees on wild bees in three areas: (1) competition for floral and nesting resources, (2) indirect effects via changes in plant communities, including the spread of exotic plants and decline of native plants, and (3) transmission of pathogens. The majority of reviewed studies reported negative effects of managed bees, but trends differed across topical areas. Of studies examining competition, results were highly variable with 53% reporting negative effects on wild bees, while 28% reported no effects and 19% reported mixed effects (varying with the bee species or variables examined). Equal numbers of studies examining plant communities reported positive (36%) and negative (36%) effects, with the remainder reporting no or mixed effects. Finally, the majority of studies on pathogen transmission (70%) reported potential negative effects of managed bees on wild bees. However, most studies across all topical areas documented the potential for impact (e.g. reporting the occurrence of competition or pathogens), but did not measure direct effects on wild bee fitness, abundance, or diversity. Furthermore, we found that results varied depending on whether managed bees were in their native or non-native range; managed bees within their native range had lesser competitive effects, but potentially greater effects on wild bees via pathogen transmission. We conclude that while this field has expanded considerably in recent decades, additional research measuring direct, long-term, and population-level effects of managed bees is needed to understand their potential impact on wild bees.
By facilitating plant reproduction, pollinators perform a crucial ecological function that supports the majority of the world's plant diversity, and associated organisms, and a significant fraction of global agriculture. Thus, pollinators are simultaneously vital to supporting both natural ecosystems and human food security, which is a unique position for such a diverse group of organisms. The past two decades have seen unprecedented interest in pollinators and pollination ecology, stimulated in part by concerns about the decline of pollinator abundance and diversity in some parts of the world. This review synthesizes what is currently understood about the taxonomic diversity of organisms that are known to act as pollinators; their distribution in both deep time and present space; the importance of their diversity for ecological function (including agro-ecology); changes to diversity and abundance over more recent timescales, including introduction of non-native species; and a discussion of arguments for conserving their diversity.
Honey bees are important pollinators of agricultural crops. Pathogens and other factors have been implicated in high annual losses of honey bee colonies in North America and some European countries. To further investigate the relationship between multiple factors, including pathogen prevalence and abundance and colony health, we monitored commercially managed migratory honey bee colonies involved in California almond pollination in 2014. At each sampling event, honey bee colony health was assessed, using colony population size as a proxy for health, and the prevalence and abundance of seven honey bee pathogens was evaluated using PCR and quantitative PCR, respectively. In this sample cohort, pathogen prevalence and abundance did not correlate with colony health, but did correlate with the date of sampling. In general, pathogen prevalence (i.e., the number of specific pathogens harbored within a colony) was lower early in the year (January—March) and was greater in the summer, with peak prevalence occurring in June. Pathogen abundance in individual honey bee colonies varied throughout the year and was strongly associated with the sampling date, and was influenced by beekeeping operation, colony health, and mite infestation level. Together, data from this and other observational cohort studies that monitor individual honey bee colonies and precisely account for sampling date (i.e., day of year) will lead to a better understanding of the influence of pathogens on colony mortality and the effects of other factors on these associations.
Basil (Ocimum L.) species are used as medicinal plants due to their essential oils exhibiting specific biological activity. The present work demonstrated that both the variety and season/conditions of cultivation had a significant effect on (i) the produced amount (extraction yield), (ii) qualitative, as well as (iii) quantitative profile of basil essential oil. Among studied basil varieties, a new variety, ‘Mánes’, was characterized for the first time. Based on our quantitative evaluation of GC-MS profiles, the following chemotypes and average concentrations of a main component were detected in the studied basil varieties: ‘Ohře’, ‘Lettuce Leaf’, ‘Purple Opaal’, ‘Dark Green’ (linalool, 5.99, 2.49, 2.34, 2.01 mg/mL, respectively), and ‘Mammolo Genovese’, ‘Mánes’, ‘Red Rubin’ (eucalyptol, 1.34, 0.96, 0.76 mg/mL, respectively). At the same time, when considering other compounds identified in GC-MS profiles, all the studied varieties, except from ‘Lettuce Leaf’, were methyl eugenol-rich with a strong dependence of the eugenol:methyl eugenol ratio on the seasonal changes (mainly solar irradiation, but also temperature and relative humidity). More complex and/or variable (depending on the season and cultivation) chemotypes were observed with ‘Lettuce Leaf’ (plus estragole, 2.27 mg/mL), ‘Dark Green’ (plus eucalyptol, 1.36 mg/mL), ‘Mammolo Genovese’ (plus eugenol, 1.19 mg/mL), ‘Red Rubin’ (plus linalool and eugenol, 0.46 and 0.56 mg/mL, respectively), and ‘Mánes’ (plus linalool and eugenol, 0.58 and 0.40 mg/mL, respectively). When considering superior extraction yield (ca. 17 mL·kg−1, i.e., two to five times higher than other examined varieties) and consistent amounts (yields) of essential oil when comparing inter-seasonal or inter-year data (RSD and inter-year difference in mean yield values ˂2.5%), this new basil variety is very promising for use in the pharmaceutical, food, and cosmetic industries.
Pollination is a valuable ecosystem service, and plant–pollinator interactions in particular are known to play a crucial role in conservation and ecosystem functioning. These mutualisms, like other ecological interactions, are currently threatened by different drivers of global change, mainly habitat loss, fragmentation, or modification of its quality. Most studies so far have focused on the impact of such disturbances on particular species interactions and we thus need more empirical evidence on the responses at a community-level. Here we evaluated how habitat loss influenced the pattern of interactions between plants and their flower visitors in a coastal dune marshland community. Using data from four years (2008–2011), we assessed the effect of a large disturbance in the area (occurring in 2010) that represented the loss of more than 50% of the vegetation cover. We found a considerable decrease in species richness and abundance of flower visitors, which resulted in a lower number of interactions after the disturbance. Not all functional groups, however, responded similarly. Contrary to the expected from previous findings, bees and wasps were less negatively influenced than beetles, flies and ants, possibly due to their higher movement capacity. Species interactions in the community were more specialized after habitat loss, resulting in a lower level of network nestedness and a higher modularity. At a species level, the number of flower visitors per plant decreased after the disturbance, and plants were visited by less abundant flower visitors. Our findings lead us to predict that the overall plant–flower visitor network became less robust and resilient to future perturbations. However, the fact that each functional group responds distinctly to disturbances makes it more difficult to foresee the final consequences on community composition and ecosystem functioning.
Damage confirmed
Early studies of the impacts of neonicotinoid insecticides on insect pollinators indicated considerable harm. However, lingering criticism was that the studies did not represent field-realistic levels of the chemicals or prevailing environmental conditions. Two studies, conducted on different crops and on two continents, now substantiate that neonicotinoids diminish bee health (see the Perspective by Kerr). Tsvetkov et al. find that bees near corn crops are exposed to neonicotinoids for 3 to 4 months via nontarget pollen, resulting in decreased survival and immune responses, especially when coexposed to a commonly used agrochemical fungicide. Woodcock et al. , in a multicounty experiment on rapeseed in Europe, find that neonicotinoid exposure from several nontarget sources reduces overwintering success and colony reproduction in both honeybees and wild bees. These field results confirm that neonicotinoids negatively affect pollinator health under realistic agricultural conditions.
Science , this issue p. 1395 , p. 1393 ; see also p. 1331
Many plants defend themselves against herbivores by chemical deterrents in their tissues and the presence of such substances in floral nectar means that pollinators often encounter them when foraging. The effect of such substances on the foraging behaviour of pollinators is poorly understood. Using artificial flowers in tightly-controlled laboratory settings, we examined the effects of the alkaloid nicotine on bumblebee foraging performance. We found that bumblebees confronted simultaneously with two equally rewarded nicotine-containing and nicotine-free flower types are deterred only by unnaturally high nicotine concentrations. This deterrence disappears or even turns into attraction at lower nectar-relevant concentrations. The alkaloid has profound effects on learning in a dose-dependent manner. At a high natural dose, bees learn the colour of a nicotine-containing flower type more swiftly than a flower type with the same caloric value but without nicotine. Furthermore, after experiencing flowers containing nicotine in any tested concentration, increasing numbers of bumblebees stay more faithful to these flowers, even if they become a suboptimal choice in terms of reward. These results demonstrate that alkaloids enhance pollinator flower constancy, opening new perspectives in co-evolutionary process between plants and pollinators.
During the mid-Cretaceous, angiosperms diversified from several nondiverse lineages to their current global domination [1], replacing earlier gymnosperm lineages [2].Several hypotheses explain this extensive radiation [3], one of which involves proliferation of insect pollinator associations in the transition from gymnosperm to angiosperm dominance. However, most evidence supports gymnosperm–insect pollinator associations, buttressed by direct evidence of pollen on insect bodies, currently established for four groups: Thysanoptera (thrips), Neuroptera (lacewings), Diptera (flies), and now Coleoptera (beetles). Each group represents a distinctive pollination mode linked to a unique mouthpart type and feeding guild [4–9]. Extensive indirect evidence, based on specialized head and mouthpart morphology, is present for one of these pollinator types, the long-proboscid pollination mode [10], representing minimally ten family-level lineages of Neuroptera, Mecoptera (scorpionflies), and Diptera [8, 10, 11]. A recurring feature uniting these pollinator modes is host associations with ginkgoalean, cycad, conifer, and bennettitalean gymnosperms. Pollinator lineages bearing these pollination modes were categorized into four evolutionary cohorts during the 35-million year-long angiosperm radiation, each defined by its host-plant associations (gymnosperm or angiosperm) and evolutionary pattern (extinction, continuation, or origination) during this interval [12]. Here, we provide the first direct evidence for one cohort, exemplified by the beetle Darwinylus marcosi, family Oedemeridae (false blíster beetles), that had an earlier gymnosperm (most likely cycad) host association, later transitioning onto angiosperms [13]. This association constitutes one of four patterns explaining the plateau of family-level plant lineages generally and pollinating insects specifically during the mid-Cretaceous angiosperm radiation [12].
Although bee-plant associations are generally maintained through speciation processes, host shifts have occurred during evolution. Understanding shifts between both phylogenetically and morphologically unrelated plants (i.e., host-saltation) is especially important since they could have been key processes in the origin and radiation of bees. Probably far from being a random process, such host-saltation might be driven by hidden constraints associated with plant traits. We selected two clades of oligolectic bees (i.e., Colletes succinctus group and Melitta leporina group) foraging on co-flowering but unrelated host-plants to test this hypothesis. We analyzed floral scent, floral color and chemical composition of pollen from host and non-host plants of these two clades. We did not find evidence for host-plant evolution in the Melitta leporina group driven by one of the assayed floral traits. On the contrary, hosts of the C. succinctus group display similar primary nutritive content of pollen (i.e., amino acids and sterols) but not similar floral scent or color, suggesting that shared pollen chemistry probably mediates saltation in this clade. Our study revealed that constraints shaping floral associations are diverse and clearly depend on species life-history traits, but evidence suggests that pollen chemistry may act as a major floral filter and guide evolutionary host-shifts.
For millennia, humans have imposed strong selection on domesticated crops, resulting in drastically altered crop phenotypes compared with wild ancestors. Crop yields have increased, but a long-held hypothesis is that domestication has also unintentionally decreased plant defences against herbivores. To test this hypothesis, we conducted a phylogenetically controlled meta-analysis comparing insect herbivore resistance and putative plant defence traits between crops and their wild relatives. Our database included 2098 comparisons made across 73 crops in 89 studies. We found that domestication consistently reduced plant resistance to herbivores, although the magnitude of the effects varied across plant organs and depended on how resistance was measured. However, domestication had no consistent effects on the specific plant defence traits underlying resistance, including secondary metabolites and physical feeding barriers. The values of these traits sometimes increased and sometimes decreased during domestication. Consistent negative effects of domestication were observed only when defence traits were measured in reproductive organs or in the plant organ that was harvested. These results highlight the complexity of evolution under domestication and the need for an improved theoretical understanding of the mechanisms through which agronomic selection can influence the species interactions that impact both the yield and sustainability of our food systems.
This article is part of the themed issue ‘Human influences on evolution, and the ecological and societal consequences’.
Nectar and pollen contain diverse phytochemicals that can reduce disease in pollinators. However, prior studies showed variable effects of nectar chemicals on infection, which could reflect variable phytochemical resistance among parasite strains. Inter-strain variation in resistance could influence evolutionary interactions between plants, pollinators, and pollinator disease, but testing direct effects of phytochemicals on parasites requires elimination of variation between bees. Using cell cultures of the bumble bee parasite Crithidia bombi, we determined (1) growth-inhibiting effects of nine floral phytochemicals and (2) variation in phytochemical resistance among four parasite strains. C. bombi growth was unaffected by naturally occurring concentrations of the known antitrypanosomal phenolics gallic acid, caffeic acid, and chlorogenic acid. However, C. bombi growth was inhibited by anabasine, eugenol, and thymol. Strains varied >3-fold in phytochemical resistance, suggesting that selection for phytochemical resistance could drive parasite evolution. Inhibitory concentrations of thymol (4.53–22.2 ppm) were similar to concentrations in Thymus vulgaris nectar (mean 5.2 ppm). Exposure of C. bombi to naturally occurring levels of phytochemicals—either within bees or during parasite transmission via flowers—could influence infection in nature. Flowers that produce antiparasitic phytochemicals, including thymol, could potentially reduce infection in Bombus populations, thereby counteracting a possible contributor to pollinator decline.
Factors associated with agricultural intensification, for example, loss of seminatural vegetation and pesticide use has been shown to adversely affect the bee community. These factors may impact the bee community differently at different landscape scales. The scale dependency is expected to be more pronounced in heterogeneous landscapes. However, the scale-dependent response of the bee community to drivers of its decline is relatively understudied, especially in the tropics where the agricultural landscape is often heterogeneous. This study looked at effects of agricultural intensification on bee diversity at patch and landscape scales in a tropical agricultural landscape. Wild bees were sampled using 12 permanent pan trap stations. Patch and landscape characteristics were measured within a 100 m (patch scale) and a 500 m (landscape scale) radius of pan trap stations. Information on pesticide input was obtained from farmer surveys. Data on vegetation cover, productivity, and percentage of agricultural and fallow land (FL) were collected using satellite imagery. Intensive areas in a bee-site network were less specialized in terms of resources to attract rare bee species while the less intensive areas, which supported more rare species, were more vulnerable to disturbance. A combination of patch quality and diversity as well as pesticide use regulates species diversity at the landscape scale (500 m), whereas pesticide quantity drove diversity at the patch scale (100 m). At the landscape scale, specialization of each site in terms of resources for bees increased with increasing patch diversity and FL while at the patch scale specialization declined with increased pesticide use. Bee functional groups responded differentially to landscape characteristics as well as pesticide use. Wood nesting bees were negatively affected by the number of pesticides used but other bee functional groups were not sensitive to pesticides. Synthesis and Applications: Different factors affect wild bee diversity at the scale of landscape and patch in heterogeneous tropical agricultural systems. The differential response of bee functional groups to agricultural intensification underpins the need for guild-specific management strategies for wild bee conservation. Less intensively farmed areas support more rare species and are vulnerable to disturbance; consequently, these areas should be prioritized for conservation to maintain heterogeneity in the landscape. It is important to conserve and restore seminatural habitats to maintain complexity in the landscapes through participatory processes and to regulate synthetic chemical pesticides in farm operations to conserve the species and functional diversity of wild bees.
The reproductive organs and mating biology of angiosperms exhibit greater variety than those of any other group of organisms. Flowers and inflorescences are also the most diverse structures produced by angiosperms, and floral traits provide some of the most compelling examples of evolution by natural selection. Given that flowering plants include roughly 250,000 species, their reproductive diversity will not be explained easily by continued accumulation of case studies of individual species. ınstead a more strategic approach is now required, which seeks to identify general principles concerning the role of ecological function in the evolution of reproductive diversity. The Ecology and Evolution of Flowers uses this approach to expose new insights into the functional basis of floral diversity, and presents the very latest theoretical and empirical research on floral evolution. Floral biology is a dynamic and growing area and this book, written by the leading internationally recognized researchers in this field, reviews current progress in understanding the evolution and function of flowers. Chapters contain both new research findings and synthesis. Major sections in turn examine functional aspects of floral traits and sexual systems, the ecological influences on reproductive adaptation, and the role of floral biology in angiosperm diversification. Overall, this integrated treatment illustrates the role of floral function and evolution in the generation of angiosperm biodiversity. This advanced textbook is suitable for graduate level students taking courses in plant ecology, evolution, systematics, biodiversity and conservation. ıt will also be of interest and use to a broader audience of plant scientists seeking an authoritative overview of recent advances in floral biology.
Global pollinator declines and land-use change can lead to pollination limitation with implications for agricultural productivity. Hand pollination is used in agricultural production as a technique to manually pollinate crops. But the prevalence of hand pollination, as well as benefits and costs, remain unknown. We systematically reviewed the literature for examples, methods, drivers, and economic motivations of hand pollination. Furthermore, we discuss the risks, constraints, and opportunities of hand pollination. We found evidence for 20 hand-pollinated crops, including minor but also economically important crops (e.g. apple, oil palm, cacao). The lack of pollinators was the most important reason for the application of hand pollination (50% of crops), while insufficient proportion or proximity of pollinizers (8% of crops) and skewed sex ratio or dichogamy (8% of crops) were second most important. The main economic motivations for practicing or recommending hand pollination were to increase fruit set, and/or fruit quality (78% of crops). Hand pollination is practiced in large- and small-scale farming, home gardens, and greenhouses. Opportunities of hand pollination are the control of pollen origin and quantity, pollination timing and frequency as well as independence from environmental fluctuations. Farmers can increase yields, improve fruit quality, avoid fruit abortion, increase employment, and secure subsistence food. The main constraints of hand pollination are high labor inputs, high material costs, and required skills. Major risks of hand pollination include management ignoring pollinator conservation, high food prices, over-pollination, labor accidents, and unfair labor. We conclude that in the face of global change, hand pollination allows improved control of pollination and is likely to increase in importance. The benefits of hand pollination need to outweigh the costs and fair labor is essential. Altogether, hand pollination can be a valuable tool for crop systems where pollinators are absent or are not reliable for sustaining high-quality crop production.
The phrase "Darwin's abominable mystery" is frequently used with reference to a range of outstanding questions about the evolution of the plant group today known as the angiosperms. Here, I seek to more fully understand what prompted Darwin to coin the phrase in 1879, and the meaning he attached to it, by surveying the systematics, paleobotanical records, and phylogenetic hypotheses of his time. In the light of this historical research, I argue that Darwin was referring to the origin only of a subset of what are today called angiosperms: a (now obsolete) group equivalent to the "dicotyledons" of the Hooker and Bentham system. To Darwin and his contemporaries, the dicotyledons' fossil record began abruptly and with great diversity in the Cretaceous, whereas the gymnosperms and monocotyledons were thought to have fossil records dating back to the Carboniferous or beyond. Based on their morphology, the dicotyledons were widely seen by botanists in Darwin's time (unlike today) as more similar to the gymnosperms than to the monocotyledons. Thus, morphology seemed to point to gymnosperm progenitors of dicotyledons, but this hypothesis made the monocotyledons, given their (at the time) apparently longer fossil record, difficult to place. Darwin had friendly disagreements about the mystery of the dicotyledons' abrupt appearance in the fossil record with others who thought that their evolution must have been more rapid than his own gradualism would allow. But the mystery may have been made "abominable" to him because it was seen by some contemporary paleobotanists, most notably William Carruthers, the Keeper of Botany at the British Museum, as evidence for divine intervention in the history of life. Subsequent developments in plant systematics and paleobotany after 1879 meant that Darwin's letter was widely understood to be referring to the abrupt appearance of all angiosperms when it was published in 1903, a meaning that has been attached to it ever since.
Insect declines are being reported worldwide for flying, ground, and aquatic lineages. Most reports come from western and northern Europe, where the insect fauna is well-studied and there are considerable demographic data for many taxonomically disparate lineages. Additional cases of faunal losses have been noted from Asia, North America, the Arctic, the Neotropics, and elsewhere. While this review addresses both species loss and population declines, its emphasis is on the latter. Declines of abundant species can be especially worrisome, given that they anchor trophic interactions and shoulder many of essential ecosystem services of their respective communities. A review of the factors believed to be responsible for observed collapses and those perceived to be especially threatening to insects form the core of this treatment. In addition to widely recognized threats to insect biodiversity, e.g., habitat destruction, agricultural intensification (including pesticide use), climate change, and invasive species, this assessment highlights a few less commonly considered factors such as atmospheric nitrification from the burning of fossil fuels and the effects of droughts and changing precipitation patterns. Because the geographic extent and magnitude of insect declines are largely unknown, there is an urgent need for monitoring efforts, especially across ecological gradients, which will help to identify important causal factors in declines. This review also considers the status of vertebrate insectivores, reporting bias, challenges inherent in collecting and interpreting insect demographic data, and cases of increasing insect abundance.
Expected final online publication date for the Annual Review of Entomology, Volume 65 is January 7, 2020. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.
Beetle pollination occurs throughout the N Queensland rainforest structure, from ground-storey plants to upper canopy trees and also in pioneer species. In some pollination systems, the nectar/pollen-eating beetles predominate, in others the food-body eaters or herbivore beetles are dominant. Pollination associations range from flexible systems where cantharophily varies in importance, to specialized entirely cantharophilous systems. In the latter >1 species of beetle may be involved or the system may be so specialized, that only 1 species of beetle is the pollinator. Implications of these pollination systems for management and conservation are discussed. -from Authors
Animal pollination is an important input to the global food system, affecting 2/3 of crops and worth more than 500-137,000 per year). Conserving wild pollinator communities, therefore, can increase crop quantity, quality, and farm revenue, but some farmers will benefit more than others. Farm-specific studies and recommendations are needed to best inform local and regional management decisions.
Cycads, unlike modern wind-pollinated conifers and Ginkgo, are unusual in that they are an ancient group of gymnosperms pollinated by insects [1, 2, 3]. Although it is well documented that cycads were diverse and abundant during the mid-Mesozoic, little is known about their biogeography and pollination before the rise of angiosperms. Direct fossil evidence illuminating the evolutionary history of cycads is extremely rare [4, 5]. Here we report a specialized beetle-mediated pollination mode from the mid-Cretaceous of Myanmar, wherein a new boganiid beetle, Cretoparacucujus cycadophilus, with specialized pollen-feeding adaptations in its mouthparts and legs, was associated with many pollen grains of Cycadopites. Phylogenetic analyses indicate Cretoparacucujus as a sister group to the extant Australian Paracucujus, which pollinate the cycad Macrozamia riedlei. Our discovery, along with the current disjunct distribution of related beetle-herbivore (tribe Paracucujini) and cycad-host (tribe Encephalarteae) pairs in South Africa and Australia, indicate a probable ancient origin of beetle pollination of cycads at least in the Early Jurassic, long before angiosperm dominance and the radiation of flowering-plant pollinators later in the Cretaceous.
Current research, management and outreach programmes relevant to insect pollinator conservation are strongly focused on relationships between pollinators and insect‐pollinated crops and wild plants.
Pollinators also visit wind‐pollinated plants to collect pollen, or for nest sites and materials, but these interactions are largely overlooked. I review documented records of bee and syrphid fly species collecting pollen from wind‐pollinated plant taxa, including economically important crops, and provide the most comprehensive collation of peer‐reviewed records of pollinators visiting wind‐pollinated plants to date. I argue for more basic research into functional relationships between insect pollinators and wind‐pollinated plants.
I found over 200 visitation records for 101 wind‐pollinated plant genera in 25 families, including 4 of the 12 gymnosperm families. Almost half the records (49%) were for grasses and sedges (Poales). I also identified records of bees and/or syrphid flies visiting 10 economically important wind‐pollinated crop plant species, including three major grain crops (rice, corn, and sorghum). Most records (70%) were from indirect pollen analysis from hives, nest cells or insect bodies, highlighting the need for more direct observational studies of plant–pollinator interactions.
Insect pollinator communities require resource diversity to persist in a landscape. Hence, researchers and land managers aiming to identify links between pollinators and ecosystem function should also consider broader interactions beyond the standard traits of the entomophily syndrome.
We briefly review current understanding of wild pollinators and pollination services on farmlands.
We consider how concepts in plant ecology – community assembly and functional trait diversity ‐ may be applied to create diverse, wild pollinator communities across scales in agroecosystems.
We also make recommendations for best practices to enhance pollination services and create more sustainable food production systems under changing environmental conditions, including creating greater landscape connectivity, embracing pollinator dynamics, and providing incentives and other motivations to support these practices.
Synthesis . We highlight the opportunity for agricultural lands to serve a dual role for both food production and pollinator conservation, and conclude by posing unanswered questions and top priorities for future studies.
Estimates of inbreeding depression obtained from the literature were used to evaluate the association between inbreeding depression and the degree of self-fertilization in natural plant populations. Theoretical models predict that the magnitude of inbreeding depression will decrease with inbreeding as deleterious recessive alleles are expressed and purged through selection. If selection acts differentially among life history stages and deleterious effects are uncorrelated among stages, then the timing of inbreeding depression may also evolve with inbreeding. Estimates of cumulative inbreeding depression and stage-specific inbreeding depression (four stages: seed production of parent, germination, juvenile survival, and growth/reproduction) were compiled for 79 populations (using means of replicates, N = 62) comprising 54 species from 23 families of vascular plants. Where available, data on the mating system also were collected and used as a measure of inbreeding history. A significant negative correlation was found between cumulative inbreeding depression and the primary selfing rate for the combined sample of angiosperms (N = 35) and gymnosperms (N = 9); the correlation was significant for angiosperms but not gymnosperms examined separately. The average inbreeding depression in predominantly selfing species (δ = 0.23) was significantly less (43%) than that in predominantly outcrossing species (δ = 0.53). These results support the theoretical prediction that selfing reduces the magnitude of inbreeding depression. Most self-fertilizing species expressed the majority of their inbreeding depression late in the life cycle, at the stage of growth/reproduction (14 of 18 species), whereas outcrossing species expressed much of their inbreeding depression either early, at seed production (17 of 40 species), or late (19 species). For species with four life stages examined, selfing and outcrossing species differed in the magnitude of inbreeding depression at the stage of seed production (selfing δ = 0.05, N = 11; outcrossing δ = 0.32, N = 31), germination (selfing δ = 0.02, outcrossing δ = 0.12), and survival to reproduction (selfing δ = 0.04, outcrossing δ = 0.15), but not at growth and reproduction (selfing δ = 0.21, outcrossing δ = 0.27); inbreeding depression in selfers relative to outcrossers increased from early to late life stages. These results support the hypothesis that most early acting inbreeding depression is due to recessive lethals and can be purged through inbreeding, whereas much of the late-acting inbreeding depression is due to weakly deleterious mutations and is very difficult to purge, even under extreme inbreeding.
The amounts of inbreeding depression upon selfing and of heterosis upon outcrossing determine the strength of selection on the selfing rate in a population when this evolves polygenically by small steps. Genetic models are constructed which allow inbreeding depression to change with the mean selfing rate in a population by incorporating both mutation to recessive and partially dominant lethal and sublethal alleles at many loci and mutation in quantitative characters under stabilizing selection. The models help to explain observations of high inbreeding depression (> 50%) upon selfing in primarily outcrossing populations, as well as considerable heterosis upon outcrossing in primarily selfing populations. Predominant selfing and predominant outcrossing are found to be alternative stable states of the mating system in most plant populations. Which of these stable states a species approaches depends on the history of its population structure and the magnitude of effect of genes influencing the selfing rate.
Wild and managed pollinators provide a wide range of benefits to society in terms of contributions to food security, farmer and beekeeper livelihoods, social and cultural values, as well as the maintenance of wider biodiversity and ecosystem stability. Pollinators face numerous threats, including changes in land-use and management intensity, climate change, pesticides and genetically modified crops, pollinator management and pathogens, and invasive alien species. There are well-documented declines in some wild and managed pollinators in several regions of the world. However, many effective policy and management responses can be implemented to safeguard pollinators and sustain pollination services.