Flower opening dynamics, pollen-ovule ratio, stigma receptivity and stigmatic pollen germination (in-vivo) in Chaenomeles speciosa (Sweet) Nakai.

Although Chaenomeles is widely used in horticulture, traditional Chinese medicine and landscape greening, insufficient research has hindered its breeding and seed selection. This study investigated the floral phenology, floral organ characteristics, palynology, and breeding systems of Chaenomeles speciosa (Sweet) Nakai. The floral characteristics of C. speciosa were observed both visually and stereoscopically. The microstructures of the flower organs were observed using scanning electron microscopy. Pollen stainability was determined using triphenyl tetrazolium chloride staining. Stigma receptivity was determined using the benzidine-H2O2 method and the post-artificial pollination pollen germination method. The breeding system was assessed based on the outcrossing index and pollen-ovule ratio. The flowers of C. speciosa were bisexual with a flowering period from March to April. The flowering periods of single flowers ranged from 8 to 19 d, and those of single plants lasted 18-20 d. The anthers were cylindrical, with the base attached to the filament, and were split longitudinally to release pollen. The flower had five styles, with a connate base. The ovaries had five carpels and five compartments. The inverted ovules were arranged in two rows on the placental axis. The stigma of C. speciosa was dry and had many papillary protrusions. In the early flowering stage (1-2 d of flowering), the pollen exhibited high stainability (up to 84.24%), but all stainability was lost at 7 d of flowering. Storage at - 20 °C effectively delayed pollen inactivation. The stigma receptivity of C. speciosa lasted for approximately 7 days, and the breeding system was classified as outcrossing with partial self-compatibility.

Heat and desiccation tolerances predict bee abundance under climate change.

Climate change could pose an urgent threat to pollinators, with critical ecological and economic consequences. However, for most insect pollinator species, we lack the long-term data and mechanistic evidence that are necessary to identify climate-driven declines and predict future trends. Here we document 16 years of abundance patterns for a hyper-diverse bee assemblage1 in a warming and drying region2, link bee declines with experimentally determined heat and desiccation tolerances, and use climate sensitivity models to project bee communities into the future. Aridity strongly predicted bee abundance for 71% of 665 bee populations (species × ecosystem combinations). Bee taxa that best tolerated heat and desiccation increased the most over time. Models forecasted declines for 46% of species and predicted more homogeneous communities dominated by drought-tolerant taxa, even while total bee abundance may remain unchanged. Such community reordering could reduce pollination services, because diverse bee assemblages typically maximize pollination for plant communities3. Larger-bodied bees also dominated under intermediate to high aridity, identifying body size as a valuable trait for understanding how climate-driven shifts in bee communities influence pollination4. We provide evidence that climate change directly threatens bee diversity, indicating that bee conservation efforts should account for the stress of aridity on bee physiology.

Defective pollen tube tip growth induces neo-polyploid infertility.

Genome duplication (generating polyploids) is an engine of novelty in eukaryotic evolution and a promising crop improvement tool. Yet newly formed polyploids often have low fertility. Here we report that a severe fertility-compromising defect in pollen tube tip growth arises in new polyploids of Arabidopsis arenosa. Pollen tubes of newly polyploid A. arenosa grow slowly, have aberrant anatomy and disrupted physiology, often burst prematurely, and have altered gene expression. These phenotypes recover in evolved polyploids. We also show that gametophytic (pollen tube) genotypes of two tip-growth genes under selection in natural tetraploid A. arenosa are strongly associated with pollen tube performance in the tetraploid. Our work establishes pollen tube tip growth as an important fertility challenge for neo-polyploid plants and provides insights into a naturally evolved multigenic solution.

Olfaction in the Anthropocene: NO3 negatively affects floral scent and nocturnal pollination.

There is growing concern about sensory pollutants affecting ecological communities. Anthropogenically enhanced oxidants [ozone (O3) and nitrate radicals (NO3)] rapidly degrade floral scents, potentially reducing pollinator attraction to flowers. However, the physiological and behavioral impacts on pollinators and plant fitness are unknown. Using a nocturnal flower-moth system, we found that atmospherically relevant concentrations of NO3 eliminate flower visitation by moths, and the reaction of NO3 with a subset of monoterpenes is what reduces the scent's attractiveness. Global atmospheric models of floral scent oxidation reveal that pollinators in certain urban areas may have a reduced ability to perceive and navigate to flowers. These results illustrate the impact of anthropogenic pollutants on an animal's olfactory ability and indicate that such pollutants may be critical regulators of global pollination.

Pollen, anther, stamen, and androecium mimicry.

Floral colours represent a highly diverse communication signal mainly involved in flower visitors' attraction and guidance, but also flower discrimination, filtering non-pollinators and discouraging floral antagonists. The divergent visual systems and colour preferences of flower visitors, as well as the necessity of cues for flower detection and discrimination, foster the diversity of floral colours and colour patterns. Despite the bewildering diversity of floral colour patterns, a recurrent component is a yellow UV-absorbing floral centre, and it is still not clear why this pattern is so frequent in angiosperms. The pollen, anther, stamen, and androecium mimicry (PASAM) hypothesis suggests that the system composed of the flowers possessing such yellow UV-absorbing floral reproductive structures, the flowers displaying central yellow UV-absorbing structures as floral guides, and the pollen-collecting, as well as pollen-eating, flower visitors responding to such signals constitute the world's most speciose mimicry system. In this review, we call the attention of researchers to some hypothetical PASAM systems around the globe, presenting some fascinating examples that illustrate their huge diversity. We will also present new and published data on pollen-eating and pollen-collecting pollinators' responses to PASAM structures supporting the PASAM hypothesis and will discuss how widespread these systems are around the globe. Ultimately, our goal is to promote the idea that PASAM is a plausible first approach to understanding floral colour patterns in angiosperms.

Ozone alters the chemical signal required for plant - insect pollination: The case of the Mediterranean fig tree and its specific pollinator.

Tropospheric ozone (O3) is likely to affect the chemical signal emitted by flowers to attract their pollinators through its effects on the emission of volatile organic compounds (VOCs) and its high reactivity with these compounds in the atmosphere. We investigated these possible effects using a plant-pollinator interaction where the VOCs responsible for pollinator attraction are known and which is commonly exposed to high O3 concentration episodes: the Mediterranean fig tree (Ficus carica) and its unique pollinator, the fig wasp (Blastophaga psenes). In controlled conditions, we exposed fig trees bearing receptive figs to a high-O3 episode (5 h) of 200 ppb and analyzed VOC emission. In addition, we investigated the chemical reactions occurring in the atmosphere between O3 and pollinator-attractive VOCs using real-time monitoring. Finally, we tested the response of fig wasps to the chemical signal when exposed to increasing O3 mixing ratios (0, 40, 80, 120 and 200 ppb). The exposure of the fig tree to high O3 levels induced a significant decrease in leaf stomatal conductance, a limited change in the emission by receptive figs of VOCs not involved in pollinator attraction, but a major change in the relative abundances of the compounds among pollinator-attractive VOCs in O3-enriched atmosphere. Fig VOCs reacted with O3 in the atmosphere even at the lowest level tested (40 ppb) and the resulting changes in VOC composition significantly disrupted the attraction of the specific pollinator. These results strongly suggest that current O3 episodes are probably already affecting the interaction between the fig tree and its specific pollinator.

Discussion: Harnessing microbiome-mediated adaptations in insect pollinators to mitigate climate change impact on crop pollination.

Insect pollinators, vital for agriculture and biodiversity, face escalating threats from climate change. We argue and explore the pivotal role of the microbiomes in shaping adaptations of insect pollinator resilience amid climate-induced challenges (climate change and habitat alteration). Examining diverse taxonomic groups, we unravel the interplay between insect physiology, microbiomes, and adaptive mechanisms. Climate-driven alterations in microbiomes impact insect health, behavior, and plant interactions, posing significant effects on agricultural ecosystems. We propose harnessing microbiome-mediated adaptations as a strategic approach to mitigate climate change impacts on crop pollination. Insights into insect-pollinator microbiomes offer transformative avenues for sustainable agriculture, including probiotic interventions (use of EM PROBIOTIC) and microbiome engineering (such as engineering gut bacteria) to induce immune responses and enhanced pollination services. Integrating microbiome insights into conservation practices elucidates strategies for preserving pollinator habitats, optimizing agricultural landscapes, and developing policies to safeguard pollinator health in the face of environmental changes. Finally, we stress interdisciplinary collaboration and the urgency of understanding pollinator microbiome dynamics under climate change in future research.

Adaptive foraging of pollinators fosters gradual tipping under resource competition and rapid environmental change.

Plant and pollinator communities are vital for transnational food chains. Like many natural systems, they are affected by global change: rapidly deteriorating conditions threaten their numbers. Previous theoretical studies identified the potential for community-wide collapse above critical levels of environmental stressors-so-called bifurcation-induced tipping points. Fortunately, even as conditions deteriorate, individuals have some adaptive capacity, potentially increasing the boundary for a safe operating space where changes in ecological processes are reversible. Our study considers this adaptive capacity of pollinators to resource availability and identifies a new threat to disturbed pollinator communities. We model the adaptive foraging of pollinators in changing environments. Pollinator's adaptive foraging alters the dynamical responses of species, to the advantage of some-typically generalists-and the disadvantage of others, with systematic non-linear and non-monotonic effects on the abundance of particular species. We show that, in addition to the extent of environmental stress, the pace of change of environmental stress can also lead to the early collapse of both adaptive and nonadaptive pollinator communities. Specifically, perturbed communities exhibit rate-induced tipping points at stress levels within the safe boundary defined for constant stressors. With adaptive foraging, tipping is a more asynchronous collapse of species compared to nonadaptive pollinator communities, meaning that not all pollinator species reach a tipping event simultaneously. These results suggest that it is essential to consider the adaptive capacity of pollinator communities for monitoring and conservation. Both the extent and the rate of stress change relative to the ability of communities to recover are critical environmental boundaries.

Using individual-based trait frequency distributions to forecast plant-pollinator network responses to environmental change.

Determining how and why organisms interact is fundamental to understanding ecosystem responses to future environmental change. To assess the impact on plant-pollinator interactions, recent studies have examined how the effects of environmental change on individual interactions accumulate to generate species-level responses. Here, we review recent developments in using plant-pollinator networks of interacting individuals along with their functional traits, where individuals are nested within species nodes. We highlight how these individual-level, trait-based networks connect intraspecific trait variation (as frequency distributions of multiple traits) with dynamic responses within plant-pollinator communities. This approach can better explain interaction plasticity, and changes to interaction probabilities and network structure over spatiotemporal or other environmental gradients. We argue that only through appreciating such trait-based interaction plasticity can we accurately forecast the potential vulnerability of interactions to future environmental change. We follow this with general guidance on how future studies can collect and analyse high-resolution interaction and trait data, with the hope of improving predictions of future plant-pollinator network responses for targeted and effective conservation.

Embryological observations on seed abortion in Hibiscus syriacus L. and physiological studies on nutrients, enzyme activity and endogenous hormones.

Under natural conditions, most Hibiscus syriacus L. individuals form very few mature seeds or the mature seeds that do form are of poor quality. As a result, seed yield is poor and seeds have low natural germinability. These phenomena strongly hinder utilization of the excellent germplasm resources of H. syriacus. The study has shown that pollen activity and stigma receptivity were high on the day of anthesis, and the pistils and stamens were fertile. Pollen release and stigma receptivity were synchronous. But in styles following self and cross-pollination, pollen tube abnormalities (distortion and twisting of the pollen tubes) and callose deposition were observed. Cross-pollinated pollen tubes elongated faster and fewer pollen tube abnormalities were observed compared with self-pollinated pollen tubes. And during embryo development, abnormalities during the heart-shaped embryo stage led to embryo abortion. Imbalance in antioxidant enzyme activities and low contents of auxin and cytokinin during early stages of embryo development may affect embryo development. Therefore, a low frequency of outcrossing and mid-development embryo abortion may be important developmental causes of H. syriacus seed abortion. Nutrient deficiencies, imbalance in antioxidant enzyme activities, and a high content of abscisic acid at advanced stages of seed development may be physiological causes of seed abortion.

Ancient insect vision tuned for flight among rocks and plants underpins natural flower colour diversity.

Understanding the origins of flower colour signalling to pollinators is fundamental to evolutionary biology and ecology. Flower colour evolves under pressure from visual systems of pollinators, like birds and insects, to establish global signatures among flowers with similar pollinators. However, an understanding of the ancient origins of this relationship remains elusive. Here, we employ computer simulations to generate artificial flower backgrounds assembled from real material sample spectra of rocks, leaves and dead plant materials, against which to test flowers' visibility to birds and bees. Our results indicate how flower colours differ from their backgrounds in strength, and the distributions of salient reflectance features when perceived by these key pollinators, to reveal the possible origins of their colours. Since Hymenopteran visual perception evolved before flowers, the terrestrial chromatic context for its evolution to facilitate flight and orientation consisted of rocks, leaves, sticks and bark. Flowers exploited these pre-evolved visual capacities of their visitors, in response evolving chromatic features to signal to bees, and differently to birds, against a backdrop of other natural materials. Consequently, it appears that today's flower colours may be an evolutionary response to the vision of diurnal pollinators navigating their world millennia prior to the first flowers.

Association between land use, land cover, plant genera, and pollinator abundance in mixed-use landscapes.

Pollinators are threatened by land-use and land-cover changes, with the magnitude of the threat depending on the pollinating taxa, land-use type and intensity, the amount of natural habitat remaining, and the ecosystem considered. This study aims to determine the effect of land use (protected areas, plantations, pastures), land cover (percentage of forest and open areas within buffers of different sizes), and plant genera on the relative abundance of nectivorous birds (honeyeaters), bees (native and introduced), and beetles in the mixed-use landscape of the Tasman Peninsula (Tasmania, Australia) using mixed-effect models. We found the predictor selected (through model selection based on R2) and the effect of the predictors varied depending on the pollinating taxa. The land-use predictors were selected for only the honeyeater abundance model with protected areas and plantations having substantive positive effects. Land-cover predictors were selected for the honeyeater and native bee abundance models with open land cover within 1500 m and 250 m buffers having substantive negative and positive effects on honeyeaters and native bees respectively. Bees and beetles were observed on 24 plant genera of which only native plants (and not invasive/naturalised) were positively associated with pollinating insects. Pultenaea and Leucopogon were positively associated with native bees while Leucopogon, Lissanthe, Pimelea, and Pomaderris were positively associated with introduced bees. Leptospermum was the only plant genus positively associated with beetles. Our results highlight that one size does not fit all-that is pollinator responses to different landscape characteristics vary, emphasising the importance of considering multiple habitat factors to manage and support different pollinator taxa.

Enhanced plasticity and reproductive fitness of floral and seed traits facilitate non-native species spread in mountain ecosystems.

Floral and seed traits, their relationships, and responses to abiotic constraints are considered the key determinants of the invasion success of non-native plant species. However, studies evaluating the pattern of floral and seed traits of non-native species in mountain ecosystems are lacking. In this study, we determined (a) whether the floral and seed traits of native and non-native species show similarity or dissimilarity across elevations in mountains, and (b) whether the non-native species follow different allometric patterns compared with native species. Functional variations between native and non-native species were assessed through floral and seed traits: flower count, flower display area, flower mass, specific flower area, seed count, and seed mass across an elevational gradient. Permanent plots (20 × 20 m) were laid at each 100 m elevation rise from 2000 to 4000 m a.s.l. for sampling of herbaceous plant species. The mean values of floral and seed traits such as flower display area, specific flower area, and seed count were significantly higher for non-native species compared to native species. A significant difference in trait values (flower display area, flower mass, seed count, and seed mass) between non-native species and native species was observed along the elevational gradient, except for flower count and specific flower area. The bivariate relationship revealed non-native species to exhibit a stronger relationship between flower display area ∼ flower mass, and flower display area ∼ seed mass traits than the native species. Non-native species showed enhanced reproductive ability under varying environmental conditions along an elevational gradient in mountain ecosystems. Greater flower display area and seed mass at lower elevations and a stronger overall trait-trait relationship among non-native species implied resource investment in pollinator visualization, flower mass, and seed quality over seed quantity. The study concludes that enhanced plasticity and reproductive fitness of floral and seed traits would consequently aid non-native species to adapt, become invasive, and displace native species in mountain ecosystems if the climatic barriers acting on non-native species are reduced with climate change.

The Flower Colour Influences Spontaneous Nectaring in Butterflies: a Case Study with Twenty Subtropical Butterflies.

Butterflies have a wide spectrum of colour vision, and changes in flower colour influence both the visiting and nectaring (the act of feeding on flower nectar) events of them. However, the spontaneous behavioural response of butterflies while foraging on real flowers is less characterised in wild conditions. Hence, this study intends to investigate flower colour affinity in wild butterflies in relation to nectaring frequency (NF) and nectaring duration (ND). Six distinct flower colours were used to study spontaneous nectaring behaviour in 20 species of subtropical butterflies. Both NF and ND greatly varied in the flower colours they offered. Yellow flowers were frequently imbibed by butterflies for longer durations, followed by orange, while red, pink, white and violet flowers were occasionally nectared in shorter bouts. Though butterflies have a general tendency to nectar on multiple flower colours, the Nymphalids were more biased towards nectaring on yellow flowers, but Papilionids preferred both yellow and orange, while the Pierids were likely to display an equal affinity for yellow, orange and violet flowers as their first order of preference. Even if the blooms are associated with higher nectar concentrations or a significant grade reward, the butterfly may prefer to visit different-coloured flowers instead. Flower colour choice appears to be a generalist phenomenon for butterflies, but their specialist visiting nature was also significant. Nymphalid representatives responded to a wider variety of floral colour affinities than Pierid and Papilionid species. The colour preference of butterflies aids in the identification of flowers during foraging and influences subsequent foraging decisions, which ultimately benefits pollination success. The current information will support the preservation and conservation of butterflies in their natural habitats.

The interplay between scale, pollination niche and floral attractiveness on density-dependent plant-pollinator interactions.

Pollinators mediate interspecific and intraspecific plant-plant indirect interactions (competition vs. facilitation) via density-dependent processes, potentially shaping the dynamics of plant communities. However, it is still unclear which ecological drivers regulate density-dependent patterns, including scale, pollination niches (i.e., the main pollinator functional group) and floral attractiveness to pollinators. In this study, we conducted three-year field observations in Hengduan Mountains of southwest China. By gathering data for more than 100 animal-pollinated plant species, we quantified the effect (positive vs. negative) of conspecific and heterospecific flower density on pollination at two scales: plot-level (4 m2) and site-level (100-5000 m2). Then, we investigated how pollination niches and floral attractiveness to pollinators (estimated here as average per-flower visitation rates) modulated density-dependent pollination interactions. Pollinator visitation depended on conspecific and heterospecific flower density, with rare plants subjected to interspecific competition at the plot-level and interspecific facilitation at the site-level. Such interspecific competition at the plot-level was stronger for plants pollinated by diverse insects, while interspecific facilitation at the site-level was stronger for bee-pollinated plants. Moreover, we also found stronger positive conspecific density-dependence for plants with lower floral attractiveness at the site-level, meaning that they become more frequently visited when abundant. Our study indicates that the role of pollination in maintaining rare plants and plant diversity depends on the balance of density-dependent processes in species-rich communities. We show here that such balance is modulated by scale, pollination niches and floral attractiveness to pollinators, indicating the context-dependency of diversity maintenance mechanisms.

Floral Diversity and Pollination Syndromes in Agave subgenus Manfreda.

The genus Agave is an ecological keystone of American deserts and both culturally and economically important in Mexico. Agave is a large genus of about 250 species. The radiation of Agave is marked by an initial adaptation to desert environments and then a secondary diversification of species associated with pollinator groups, such as hummingbirds and nocturnal moths. Phylogenetic analyses place Agave subgenus Manfreda, or the "herbaceous agaves," in a monophyletic clade that likely evolved in part as an adaptation to novel pollination vectors. Here, we present a morphological and observational study assessing the evolution of floral form in response to pollinator specialization within this understudied group. We found significant visitation by hummingbirds and nocturnal moths to several species within the Agave subgenus Manfreda. These observations also align with our morphological analyses of floral organs and support the evolution of distinct pollination syndromes. We found that not all floral morphology is consistent within a pollination syndrome, suggesting hidden diversity in the evolution of floral phenotypes in Agave. We also characterize the morphological variation between herbarium and live specimens, demonstrating that special consideration needs to be made when combining these types of data. This work identifies the potential for studying the functional evolution of diverse floral forms within Agave and demonstrates the need to further explore ecological and evolutionary relationships to understand pollinator influence on diversification in the genus.

Chemistry, biosynthesis and biology of floral volatiles: roles in pollination and other functions.

Covering: 2010 to 2023Floral volatiles are a chemically diverse group of plant metabolites that serve multiple functions. Their composition is shaped by environmental, ecological and evolutionary factors. This review will summarize recent advances in floral scent research from chemical, molecular and ecological perspectives. It will focus on the major chemical classes of floral volatiles, on notable new structures, and on recent discoveries regarding the biosynthesis and the regulation of volatile emission. Special attention will be devoted to the various functions of floral volatiles, not only as attractants for different types of pollinators, but also as defenses of flowers against enemies. We will also summarize recent findings on how floral volatiles are affected by abiotic stressors, such as increased temperatures and drought, and by other organisms, such as herbivores and flower-dwelling microbes. Finally, this review will indicate current research gaps, such as the very limited knowledge of the isomeric pattern of chiral compounds and its importance in interspecific interactions.

Floral nectar production: what cost to a plant?

Floral nectar production is central to plant pollination, and hence to human wellbeing. As floral nectar is essentially a solution in water of various sugars, it is likely a valuable plant resource, especially in terms of energy, with plants experiencing costs/trade-offs associated with its production or absorption and adopting mechanisms to regulate nectar in flowers. Possible costs of nectar production may also influence the evolution of nectar volume, concentration and composition, of pollination syndromes involving floral nectar, and the production of some crops. There has been frequent agreement that costs of floral nectar production are significant, but relevant evidence is scant and difficult to interpret. Convincing direct evidence comes from experimental studies that relate either enhanced nectar sugar production (through repeated nectar removal) to reduced ability to produce seeds, or increased sugar availability (through absorption of additional artificial nectar) to increased seed production. Proportions of available photosynthate allocated by plants to nectar production may also indicate nectar cost. However, such studies are rare, some do not include treatments of all (or almost all) flowers per plant, and all lack quantitative cost-benefit comparisons for nectar production. Additional circumstantial evidence of nectar cost is difficult to interpret and largely equivocal. Future research should repeat direct experimental approaches that relate reduced or enhanced nectar sugar availability for a plant with consequent ability to produce seeds. To avoid confounding effects of inter-flower resource transfer, each plant should experience a single treatment, with treatment of all or almost all flowers per plant. Resource allocation by plants, pathways used for resource transfer, and the locations of resource sources and sinks should also be investigated. Future research should also consider extension of nectar cost into other areas of biology. For example, evolutionary models of nectar production are rare but should be possible if plant fitness gains and costs associated with nectar production are expressed in the same currency, such as energy. It should then be possible to understand observed nectar production for different plant species and pollination syndromes involving floral nectar. In addition, potential economic benefits should be possible to assess if relationships between nectar production and crop value are evaluated.

The role of within-plant variation in nectar production: an experimental approach.

BACKGROUND AND AIMS: Nectar, a plant reward for pollinators, can be energetically expensive. Hence, a higher investment in nectar production can lead to reduced allocation to other vital functions and/or increased geitonogamous pollination. One possible strategy employed by plants to reduce these costs is to offer variable amounts of nectar among flowers within a plant, to manipulate pollinator behaviour. Using artificial flowers, we tested this hypothesis by examining how pollinator visitation responds to inter- and intra-plant variation in nectar production, assessing how these responses impact the energetic cost per visit. METHODS: We conducted a 2 × 2 factorial experiment using artificial flowers, with two levels of nectar investment (high and low sugar concentration) and two degrees of intra-plant variation in nectar concentration (coefficient of variation 0 and 20 %). The experimental plants were exposed to visits (number and type) from a captive Bombus impatiens colony, and we recorded the total visitation rate, distinguishing geitonogamous from exogamous visits. Additionally, we calculated two estimators of the energetic cost per visit and examined whether flowers with higher nectar concentrations (richer flowers) attracted more bumblebees. KEY RESULTS: Plants in the variable nectar production treatment (coefficient of variation 20 %) had a greater proportion of flowers visited by pollinators, with higher rates of total, geitonogamous and exogamous visitation, compared with plants with invariable nectar production. When assuming no nectar reabsorption, variable plants incurred a lower cost per visit compared with invariable plants. Moreover, highly rewarding flowers on variable plants had higher rates of pollination visits compared with flowers with few rewards. CONCLUSIONS: Intra-plant variation in nectar concentration can represent a mechanism for pollinator manipulation, enabling plants to decrease the energetic costs of the interaction while still ensuring consistent pollinator visitation. However, our findings did not provide support for the hypothesis that intra-plant variation in nectar concentration acts as a mechanism to avoid geitonogamy. Additionally, our results confirmed the hypothesis that increased visitation to variable plants is dependent on the presence of flowers with nectar concentration above the mean.

The effects of urbanization on pollinators and pollination: A meta-analysis.

Urbanization is increasing worldwide, with major impacts on biodiversity, species interactions and ecosystem functioning. Pollination is an ecosystem function vital for terrestrial ecosystems and food security; however, the processes underlying the patterns of pollinator diversity and the ecosystem services they provide in cities have seldom been quantified. Here, we perform a comprehensive meta-analysis of 133 studies examining the effects of urbanization on pollinators and pollination. Our results confirm the widespread negative impacts of urbanization on pollinator richness and abundance, with Lepidoptera being the most affected group. Furthermore, pollinator responses were found to be trait-specific, with below-ground nesting and solitary Hymenoptera, and spring flyers more severely affected by urbanization. Meanwhile, cities promote non-native pollinators, which may exacerbate conservation risks to native species. Surprisingly, despite the negative effects of urbanization on pollinator diversity, pollination service measured as seed set is enhanced in non-tropical cities likely due to abundant generalists and managed pollinators therein. We emphasize that the richness of local flowering plants could mitigate the negative impacts of urbanization on pollinator diversity. Overall, the results demonstrate the varying magnitudes of multiple moderators on urban pollinators and pollination services and could help guide conservation actions for biodiversity and ecosystem function for a sustainable future.