The Importance of Ants and High-Shade Management to Coffee Pollination and Fruit Weight in Chiapas, Mexico

Recent reports show importance of pollinators to coffee and importance of ants as pollinators or floral protectors in many systems. Arthropod and pollinator diversity, however, declines with management intensification of coffee (Coffea arabica) agroecosystems. We investigated influences of both flying pollinators and ants on coffee fruit set and fruit weight in one high-shade (high-biodiversity) and one low-shade (low-biodiversity) coffee farm in Chiapas, Mexico through exclusion experiments. Contradictory to previous reports, flying pollinators alone did not affect coffee fruit set or fruit weight. Individual fruit weights, however, were higher on branches with both ants and flying pollinators (1.78 g ± 0.312 (SE)) compared to branches without ants (1.03 ± 0.029) or branches without ants or flying pollinators (1.05 ± 0.049), but only in the high-shade site. Although the mechanisms producing higher fruit weights are unknown, we discuss how ants or ant-flying pollinator interactions under high-shade coffee management may contribute to increased fruit weight and the implications of high-shade management for both sustainable coffee production and biodiversity conservation.     

Contrasting bee foraging in response to resource scale and local habitat management

It is hypothesized that two main factors drive the foraging patterns of native and exotic species: food resource availability and habitat composition. These factors are particularly relevant for native bees and exotic honeybees, essential crop pollinators that are sensitive to floral resources and habitat management, and that have recently exhibited alarming population declines. Mechanisms driving native and exotic bee foraging patterns may critically depend on floral resource availability and habitat composition, yet the impacts of these factors on bee foraging have never been simultaneously analyzed. In a coffee producing region in southern Mexico, we investigated the influence of coffee floral resource levels and habitat management on native and exotic bee foraging. We measured the amount of flowering coffee available at multiple spatial scales within two distinct agroforestry habitat types (high-shade and low-shade coffee) and recorded visits to coffee flowers, documenting bee species, visit duration and visit frequency. We observed a significantly greater number of visits in high-shade coffee habitats than in low-shade coffee habitats for both native and exotic bees. In high-shade coffee habitats, native solitary bee and native social bee visitation decreased significantly in response to increasing floral resource availability, exhibiting a 'dilution effect' at the smallest spatial scale. In contrast, in low-shade coffee habitats, Africanized honeybees exhibited a 'concentration effect', increasing visitation significantly in response to increasing floral resource availability at the largest spatial scale. This study is the first to show that foraging patterns of native bees and exotic honeybees contrast in response to floral resource level and scale and that this response is mediated by the vegetation management of the local habitat.     

Floral organs act as environmental filters and interact with pollinators to structure the yellow monkeyflower (Mimulus guttatus) floral microbiome

Assembly of microbial communities is the result of neutral and selective processes. However, the relative importance of these processes is still debated. Microbial communities of flowers, in particular, have gained recent attention because of their potential impact to plant fitness and plant‐pollinator interactions. However, the role of selection and dispersal in the assembly of these communities remains poorly understood. Here, we evaluated the role of pollinator‐mediated dispersal on the contribution of neutral and selective processes in the assembly of floral microbiomes of the yellow monkeyflower (Mimulus guttatus). We sampled floral organs from flowers in the presence and absence of pollinators within five different serpentine seeps in CA and obtained 16S amplicon data on the epiphytic bacterial communities. Consistent with strong microenvironment selection within flowers we observed significant differences in community composition across floral organs and only a small effect of geographic distance. Pollinator exposure affected the contribution of environmental selection and depended on the rate and intimacy of interactions with flower visitors. This study provides evidence of the importance of dispersal and within‐flower heterogeneity in shaping epiphytic bacterial communities of flowers, and highlights the complex interplay between pollinator behaviour, environmental selection and additional abiotic factors in shaping the epiphytic bacterial communities of flowers.     

神农架地区典型草本群落中的昆虫访花行为比较

对神农架地区不同海拔高度(1,000–2,600 m)的典型草本群落中的昆虫访花行为进行了初步的监测和统计。在9个样地中共观察到鳞翅目、膜翅目、双翅目11科昆虫访花, 访花者的类群在不同海拔高度的群落中有所不同, 但明显与植物群落的物种组成有关。对昆虫访花频率的监测发现, 不同物种组成的草本群落中, 所有昆虫访花的总频率和不同类群的昆虫访花频率都有差异, 正在开花的物种较多的群落中通常能吸引相对较多的访花者。同一种植物在不同的群落背景中受访频率差异显著, 如花被不甚显著的酸模(Rumex acetosa)在某些群落中未观察到访花者, 在另一些群落中却有昆虫频繁访问。而花被显著的血见愁老鹳草(Geranium henryi)在各群落中都有频繁的昆虫访问。一些植物(如广布野豌豆Vicia cracca)虽然在不同群落中都有昆虫访问, 但访花者类群却有较大差异。作者认为, 在植物群落中, 不同植物的花在吸引昆虫传粉方面存在着一定的相互作用, 在研究特定植物的虫媒传粉时应重视该植物种群所处的群落背景所起的作用。     

Local adaptation: Mechanical fit between floral ecotypes of Nerine humilis (Amaryllidaceae) and pollinator communities

Geographic variation in floral morphology is often assumed to reflect geographic variation in pollinator communities and associated divergence in selective pressures. We studied populations of Nerine humilis (Amaryllidaceae) to assess whether geographic variation in floral form is the result of local adaptation to different pollinator communities. We first tested for associations between floral traits and visitor communities, and found that populations with similar floral morphologies were visited by similar insect communities. Mean style length in each population was also closely associated with the mean body length of the local visitor community. A reciprocal translocation experiment demonstrated that native phenotypes set more seed than translocated phenotypes. Single visitation experiments showed that native flowers received more pollen, and set more seed per visit, than introduced phenotypes in both populations. This suggests that the effectiveness of pollinator visits is determined by the degree of mechanical fit between flowers and visitors. We provide strong evidence that the observed among‐population variation in floral traits is an adaptive response to geographic variation in the pollinator community.     

胜山阔叶红松林蚂蚁巢穴对其他地表节肢动物多样性的影响

为了阐明蚂蚁巢穴对其他地表节肢动物群落组成和多样性的影响,于2019年在胜山自然保护区的原始阔叶红松林内开展实验,并采用陷阱法收集地表节肢动物.共捕获地表节肢动物92266只,分别为蜈蚣目、盲蛛目、马陆目、蜘蛛目,大头蚁属和步甲科、隐翅虫科、葬甲科这8个类群;蚂蚁巢穴存在及到蚂蚁巢穴的不同距离对其他地表节肢动物个体数量...     

Induced Floral and Extrafloral Nectar Production Affect Ant‐pollinator Interactions and Plant Fitness

Thousands of plant species throughout tropical and temperate zones secrete extrafloral nectar to attract ants, whose presence provides an indirect defense against herbivores. Extrafloral nectaries are located close to flowers and may modify competition between ants and pollinators. Here, we used Lima bean (Phaseolus lunatus L.) to study the plants interaction between ants and flower visitors and its consequences for plant fitness. To test these objectives, we carried out two field experiments in which we manipulated the presence of ants and nectar production via induction with jasmonic acid (JA). We then measured floral and extrafloral nectar production, the number of patrolling ants and flower visitors as well as specific plant fitness traits. Lima bean plants under JA induction produced more nectar in both extrafloral nectaries and flowers, attracted more ants and produced more flowers and seeds than non‐induced plants. Despite an increase in floral nectar in JA plants, application of this hormone had no significant effects on flower visitor attraction. Finally, ant presence did not result in a decrease in the number of visits, but our results suggest that ants could negatively affect pollination efficiency. In particular, JA‐induced plants without ants produced a greater number of seeds compared with the JA‐treated plants with ants.     

Pollinator conservation at the local scale: flower density,diversity and community structure increase flower visiting insect activity to mixed floral stands

Insect pollinators play a keystone role in terrestrial ecosystems. The parallel declines in plant and pollinator communities emphasizes that plant-pollinator interactions at the community level are highly relevant for biodiversity conservation. Here we examine relationships between plants and flower-visiting insects (anthophiles) at the scale of local floral patches. We conducted a visitation survey during the spring flowering season, a peak time for pollinator activity in the threatened Cape Floristic Region, South Africa. We tested floral density, diversity and composition as predictors of anthophile diversity (measured at the family/family group level) and visitation rates in multispecies stands of flowers. Although different anthophile groups responded differently, generally anthophile visitation rates and diversity were positively affected by floral density, diversity and community structure. Anthophiles were more abundant and diverse in areas with a high density and diversity of flowers. Plant community structure affected both the likelihood of occurrence and activity of anthophiles in the plots. The two mass flowering species examined here, Relhania fruticosa and Salvia chamaeleagnea, were strong determinants of anthophile activity, greatly increasing visitation rates, even though there was, on average, lower floral density and diversity. Our results show that anthophile activity is affected by highly localised, small-scale factors, namely the density and diversity of flowers and community structure. Important among these factors are patches of high quality habitat, high in floral abundance and diversity, both of which should be included in landscape-level plans for pollinator conservation, providing stepping stones for these insects in transformed landscapes.     

Floral volatiles play a key role in specialized ant pollination

Chemical signals emitted by plants are crucial to understand the ecology and evolution of plant–animal interactions. Scent is an important component of floral phenotype and represents a decisive communication channel between plants and floral visitors. Floral volatiles promote attraction of mutualistic pollinators and, in some cases, serve to prevent flower visitation by antagonists such as ants. Despite ant visits to flowers have been suggested to be detrimental to plant fitness, in recent years there has been a growing recognition of the positive role of ants in pollination. Nevertheless, the question of whether floral volatiles mediate mutualisms between ants and ant-pollinated plants still remains largely unexplored. Here we review the documented cases of ant pollination and investigate the chemical composition of the floral scent in the ant-pollinated plant Cytinus hypocistis. By using chemical-electrophysiological analyses and field behavioural assays, we examine the importance of olfactory cues for ants, identify compounds that stimulate antennal responses, and evaluate whether these compounds elicit behavioural responses. Our findings reveal that floral scent plays a crucial role in this mutualistic ant–flower interaction, and that only ant species that provide pollination services and not others occurring in the habitat are efficiently attracted by floral volatiles. 4-oxoisophorone, (E)-cinnamaldehyde, and (E)-cinnamyl alcohol were the most abundant compounds in Cytinus flowers, and ant antennae responded to all of them. Four ant pollinator species were significantly attracted to volatiles emitted by Cytinus inflorescences as well as to synthetic mixtures and single antennal-active compounds. The small amount of available data so far suggest that there is broad interspecific variation in floral scent composition among ant-pollinated plants, which could reflect differential responses and olfactory preferences among different ant species. Many exciting discoveries will be made as we enter into further research on chemical communication between ants and plants.     

Generalized and complementary pollination system in the Andean cactus Echinopsis schickendantzii

The considerable floral diversity present in the cactus family has often been associated with the specificity of its pollinators. However, many cactus pollination systems are generalized as their flowers are pollinated by a wide spectrum of animals. For example, cactus species with white flowers, nocturnal anthesis and extended floral cycles would present generalized pollination systems in which both nocturnal and diurnal visitors could be effective pollinators. In this article, we tested this hypothesis by studying the pollination biology of Echinopsis schickendantzii, an Andean cactus with sphingophilous flowers. In addition, we evaluated whether the cactus’s pollination system is complementary or redundant regarding the relative contributions of nocturnal and diurnal pollinators. Specifically, we studied the floral cycle, the reproductive system and the pollination effectiveness of floral visitors. The flowers of E. schickendantzii are self-incompatible; they opened at crepuscule and have an extended floral cycle. Moths were frequent visitors at night, whereas bees were frequent visitors during the day; both were effective pollinators of the cactus. Our results indicated that the flowers of this species present phenotypic, functional and ecological generalization, and their fruit set is determined by the contributions of both pollinator functional groups, i.e., they have complementary pollination systems. These results support the hypothesis that cacti in the extra-tropical deserts of South America have generalized pollination systems.     

Ant pollination of the palaeoendemic dioeciousBorderea pyrenaica (Dioscoreaceae)

The importance of ants for pollination in the dioeciousBorderea pyrenaica (Dioscoreaceae), a Tertiary palaeoendemic plant of the Pyrenees (NE Iberian Peninsula) was studied. The frequency of different visitors (ants, lady beetles, andDiptera) to staminate and pistillate flowers was quantified, and their effectiveness as pollinators was examined by means of fruit and seed set in selective experimental exclusions. Although ants were less abundant on flowers than other visitors, they were the most effective pollinators. Some qualitative factors of this mutualistic ant-plant interaction may account for their effectiveness: the small size of the ants, their high visitation rate to pistillate flowers, and the lack of reduction in viability of the pollen transported on the integument. In addition, the sedentary nature of ants assures their presence during the flowering period. The most abundant floral visitors ofB. pyrenaica were therefore not the most effective pollinators.     

Slippery flowers as a mechanism of defence against nectar-thieving ants

Background and AimsThe great diversity of floral characteristics among animal-pollinated plants is commonly understood to be the result of coevolutionary interactions between plants and pollinators. Floral antagonists, such as nectar thieves, also have the potential to exert an influence upon the selection of floral characteristics, but adaptation against floral antagonists has attracted comparatively little attention. We found that the corollas of hornet-pollinated Codonopsis lanceolata (Campanulaceae) and the tepals of bee-pollinated Fritillaria koidzumiana (Liliaceae) are slippery to nectar-thieving ants living in the plant’s habitat; because the flowers of both species have exposed nectaries, slippery perianths may function as a defence against nectar-thieving ants.MethodsWe conducted a behavioural experiment and observed perianth surface microstructure by scanning electron microscopy to investigate the mechanism of slipperiness. Field experiments were conducted to test whether slippery perianths prevent floral entry by ants, and whether ant presence inside flowers affects pollination.Key ResultsScanning electron microscopy observations indicated that the slippery surfaces were coated with epicuticular wax crystals. The perianths lost their slipperiness when wiped with hexane. Artificial bridging of the slippery surfaces using non-slippery materials allowed ants to enter flowers more frequently. Experimental introduction of live ants to the Codonopsis flowers evicted hornet pollinators and shortened the duration of pollinator visits. However, no statistical differences were found in the fruit or seed sets of flowers with and without ants.ConclusionsSlippery perianths, most probably based on epicuticular wax crystals, prevent floral entry by ants that negatively affect pollinator behaviour. Experimental evidence of floral defence based on slippery surfaces is rare, but such a mode of defence may be widespread amongst flowering plants.     

Drought reduces floral resources for pollinators

Climate change is predicted to result in increased occurrence and intensity of drought in many regions worldwide. By increasing plant physiological stress, drought is likely to affect the floral resources (flowers, nectar and pollen) that are available to pollinators. However, little is known about impacts of drought at the community level, nor whether plant community functional composition influences these impacts. To address these knowledge gaps, we investigated the impacts of drought on floral resources in calcareous grassland. Drought was simulated using rain shelters and the impacts were explored at multiple scales and on four different experimental plant communities varying in functional trait composition. First, we investigated the effects of drought on nectar production of three common wildflower species (Lathyrus pratensis, Onobrychis viciifolia and Prunella vulgaris). In the drought treatment, L. pratensis and P. vulgaris had a lower proportion of flowers containing nectar and O. viciifolia had fewer flowers per raceme. Second, we measured the effects of drought on the diversity and abundance of floral resources across plant communities. Drought reduced the abundance of floral units for all plant communities, irrespective of functional composition, and reduced floral species richness for two of the communities. Functional diversity did not confer greater resistance to drought in terms of maintaining floral resources, probably because the effects of drought were ubiquitous across component plant communities. The findings indicate that drought has a substantial impact on the availability of floral resources in calcareous grassland, which will have consequences for pollinator behaviour and populations.     

Fine-scale temporal dynamics of flower visitors sheds light on insect-assemblage overlap between sexes in a dioecious Ecuadorian palm

Dioecious plants generally display sexual dimorphism in male and female floral traits, potentially attracting slightly different pollinator communities. The sharing of common floral visitors between male and female flowers and their timing of visits to both sexes is of critical importance to ensure plant's reproductive success. Palm inflorescences are visited by abundant and diverse insect communities, yet the temporal patterns of insect visits on both sexes remain poorly known. We characterized the composition of a community of flower-visiting arthropods associated with the dioecious ivory palm (Phytelephas aequatorialis, Spruce) in a pre-montane forest of Ecuador. We monitored the temporal variations in insect visits along the flowering of 12 inflorescences (eight female and four male) using interception traps recovered every 4 h. We report 59 morphospecies in the arthropod community, dominated by three beetle families: Staphylinidae, Nitidulidae, and Curculionidae. Male inflorescences were more abundantly visited than female, but visitors of the later were taxonomically more diverse. Among the 16 pollinator candidates identified, nine visited both inflorescence sexes synchronously at dusk /night whereas the others did so asynchronously during the day. Our study provides new insights into the pollination mechanism of P. aequatorialis. We found evidence of differential pollinator attraction between floral sexes, which may be explained by the sexual dimorphism of both flowers. Synchronicity in dusk/night visits of both inflorescence sexes suggests a sexual synchronization of the signal used to attract pollinators.     

Ants climb plants because they cannot swim: ant presence on flowers during the flood season reduces the frequency of floral visitors

1. Seasonal changes in environments may not only affect habitat connectivity but may also affect its use by species and their interactions. Thus, during the flood season, ants are forced to develop survival strategies such as vertical plant migration.
2. According to this, it has been hypothesized that the presence of ants may directly affect plant-pollinator interactions.
3. Thus, we asked the following questions: (i) Are floral visitors of Hyptis brevipes expelled due to ant presence on inflorescences during the flood period? (ii) Is the ant effect mediated by the abundance of ants foraging on inflorescences? And, (iii) Does flower abundance predict the abundance of floral visits and ants?
4. We experimentally sampled 59 H. brevipes plants with and without ants during the flooded season, and observed no differences in flower abundance between ant treatments.
5. The probability of detaining floral visitors on H. brevipes increased with ant abundance and exceeded 50% possible repellency, but the probability of visitor deterrence was not related to flower abundance. Furthermore, the abundance of flowers did not predict the number of ants on H. brevipes individuals or the frequency of floral visits.
6. Consequently, ant repelling effects are pronounced when there are more ants foraging on plants. However, the ant repelling effect can be mitigated when plants flourish all year-round and exhibit higher concentrations of flowers in the dry months. Additionally, the different sexual functions of plants may present specific responses due to the explosive pollination mechanism associated with ant effects.

     

Floral fungal-bacterial community structure and co-occurrence patterns in four sympatric island plant species

Flowers’ fungal and bacterial communities can exert great impacts on host plant wellness and reproductive success—both directly and indirectly through species interactions. However, information about community structure and co-occurrence patterns in floral microbiome remains scarce. Here, using culture-independent methods, we investigated fungal and bacterial communities associated with stamens and pistils of four plant species (Scaevola taccada, Ipomoea cairica, Ipomoea pes-caprae, and Mussaenda kwangtungensis) growing together under the same environment conditions in an island located in South China. Plant species identity significantly influenced community composition of floral fungi but not bacteria. Stamen and pistil microbiomes did not differ in community composition, but differed in co-occurrence network topological features. Compared with the stamen network, pistil counterpart had fewer links between bacteria and fungi and showed more modular but less concentrated and connected structure. In addition, degree distribution of microbial network in each host species and each microhabitat (stamen or pistil) followed a significant power-law pattern. These results enhance our understanding in the assembly principles and ecological interactions of floral microbial communities.     

Characterization of Salix nigra floral insect community and activity of three native Andrena bees

Salix nigra (black willow) is a widespread tree that hosts many species of polylectic hymenopterans and oligolectic bees of the genus Andrena. The early flowering of S. nigra makes it an important nutritive resource for arthropods emerging from hibernation. However, since S. nigra is dioecious, not all insect visits will lead to successful pollination. Using both visual observation and pan‐trapping, we characterized the community of arthropods that visited S. nigra flowers and assessed differences among male and female trees as well as the chemical and visual drivers that influenced community composition across 3 years. We found that male trees consistently supported higher diversity of insects than female trees and only three insect species, all Andrena spp., consistently visited both sexes. Additionally, Andrena nigrae, which was the only insect that occurred more on female than male flowers, correlated strongly to volatile cues. This suggests that cross‐pollinators cue into specific aspects of floral scent, but diversity of floral visitors is driven strongly by visual cues of yellow male pollen. Through time, the floral activity of two Andrena species remained stable, but A. nigrae visited less in 2017 when flowers bloomed earlier than other years. When native bee emergence does not synchronize with bloom, activity appears to be diminished which could threaten species that subsist on a single host. Despite the community diversity of S. nigra flowers, its productivity depends on a small fraction of species that are not threatened by competition, but rather rapidly changing conditions that lead to host‐insect asynchrony.     

Reduced visitation to buzz‐pollinated Cyanella hyacinthoides in the presence of other pollen sources in the hyperdiverse Cape Floristic Region

Many plant species have floral morphologies that restrict access to floral resources, such as pollen or nectar, and only a subset of floral visitors can perform the handling behaviors required to extract restricted resources. Due to the time and energy required to extract resources from morphologically complex flowers, these plant species potentially compete for pollinators with co‐flowering plants that have more easily accessible resources. A widespread floral mechanism restricting access to pollen is the presence of tubular anthers that open through small pores or slits (poricidal anthers). Some bees have evolved the capacity to remove pollen from poricidal anthers using vibrations, giving rise to the phenomenon of buzz‐pollination. These bee vibrations that are produced for pollen extraction are presumably energetically costly, and to date, few studies have investigated whether buzz‐pollinated flowers may be at a disadvantage when competing for pollinators’ attention with plant species that present unrestricted pollen resources. Here, we studied Cyanella hyacinthoides (Tecophilaeaceae), a geophyte with poricidal anthers in the hyperdiverse Cape Floristic Region of South Africa, to assess how the composition and relative abundance of flowers with easily accessible pollen affect bee visitation to a buzz‐pollinated plant. We found that the number of pollinator species of C. hyacinthoides was not influenced by community composition. However, visitation rates to C. hyacinthoides were reduced when the relative abundances of flowers with more accessible resources were high. Visitation rates were strongly associated with petal color, showing that flower color is important in mediating these interactions. We conclude that buzz‐pollinated plants might be at a competitive disadvantage when many easily accessible pollen sources are available, particularly when competitor species share its floral signals.     

Functional diversity of ant-associated bacterial communities

It was found that ants significantly affect the physiological activity and functional diversity of soil microbial communities, and redistribution of biophilic elements (C and N) down through the profile occurs in anthills compared to the control soil, as well as their accumulation in the underground part of the ant nests. A high urease activity was revealed in ant nests and ants. Functional dissimilarities of bacterial communities in all studied objects were determined by the multisubstrate test.     

Changes amid constancy: Flower and leaf microbiomes along land use gradients and between bioregions

Microbial communities inhabiting above-ground parts of plants affect their host's development, fitness and function. Although studies on plant-associated microbes are of growing interest, environmental drivers of flower microbiomes in particular are poorly characterized. In this study, we investigated flower and leaf epiphytic bacterial microbiomes of Ranunculus acris and Trifolium pratense using metabarcoding of 16S ribosomal DNA in three German bioregions and along land-use intensity gradients. Our data suggests that the structures of bacterial communities clearly differed between plant species and tissue types. Also, floral bacterial communities of R. acris showed higher variability in comparison to T. pratense. Bacteria usually associated with pollinators were found solely in flower samples, while bacteria usually associated with the rhizosphere were only present in high abundances on leaves. We identified Pseudomonadaceae, Enterobacteriaceae and Sphingomonadaceae as the most abundant taxa on flowers, while Sphingomonadaceae, Methylobacteriaceae and Cytophagaceae dominated bacterial communities on leaves. We found that bacterial communities did not differ between long-distant regions. However, there was a turnover within each bioregion between short-distant locations. High land use intensity caused phylogenetically less diverse and more homogenous bacterial communities with an exception of T. pratense flowers. This was associated with a loss of rare bacterial families. Intensification of mowing affected the bacterial communities associated with leaves of T. pratense and fertilization led to more homogenous flower and leaf communities of R. acris, while grazing had no effects on the bacterial community composition. However, dominant taxa were not affected by land use intensification. Despite that, we identified indicator taxa for regularly disturbed environments in flower microbiomes. In conclusion, our study contributes to the knowledge about microbial community structures of the phyllosphere and extends the understanding of their community dynamics with respect to biogeographical separation and anthropogenic changes of the environment.