Nectar Research Articles

Availability of butterfly species (Lepidoptera) at different location of Bramhananda Keshab Chandra College campus and selection of respective nectar plant species

Availability of butterfly species (Lepidoptera) at different location of Bramhananda Keshab Chandra College campus and selection of respective nectar plant species

Glyphosate used as desiccant contaminates plant pollen and nectar of non-target plant species

Pesticide products containing glyphosate as a systemic active ingredient are some of the most extensively used herbicides worldwide. After spraying, residues have been found in nectar and pollen collected by bees foraging on treated plants. This dietary exposure to glyphosate could pose a hazard for flower-visiting animals including bees, and for the delivery of pollination services. Here, we evaluated whether glyphosate contaminates nectar and pollen of targeted crops and non-target wild plants. Oilseed rape was selected as focal crop species, and Rubus fruticosus growing in the hedgerows surrounding the crop was chosen as non-target plant species. Seven fields of oilseed rape, where a glyphosate-based product was applied, were chosen in east and southeast Ireland, and pollen and nectar were extracted from flowers sampled from the field at various intervals following glyphosate application. Pollen loads were taken from honeybees and bumblebees foraging on the crop at the same time. Glyphosate and aminomethylphosphonic acid (AMPA) residues were extracted using acidified methanol and their concentrations in the samples were determined by a validated liquid chromatography tandem mass spectrometry (LC-MS/MS) method. Glyphosate was detected in R. fruticosus nectar and pollen samples that were taken within a timeframe of two to seven days after the application on the crop as a desiccant. No glyphosate was detected when the application took place before or more than two months prior to our sampling in any of the evaluated matrices. The metabolite AMPA was not detected in any samples. To gain further insight into the potential extent of translocation within both plants and soil when a crop is desiccated using glyphosate before harvesting, and the potential impacts on bees, we recommend a longitudinal study of the presence and fate of glyphosate in non-target flowering plants growing nearby crop fields, over a period of several days after glyphosate application.

Herbivory, plant traits and nectar chemistry interact to affect the community of insect visitors and pollination in common milkweed, Asclepias syriaca.

Herbivory can alter plant fitness directly through changing reproductive allocation and indirectly through changing pollinator identity or behavior. Common milkweed is a plant of conservation concern with an inducible chemical defense that is also an important nectar resource. In this study, we aim to understand how herbivory severity and plant traits, including morphology and nectar chemistry, interact to affect insect visitation and pod production in common milkweed. We conducted pollinator watches on plants with experimentally varied herbivory severity and quantified insect frequency and visit length as a response to nectar chemistry, ramet height, number of inflorescences, number of flowers per inflorescence and percent tissue removed. We also quantified pollinator effectiveness and importance. Increased herbivory severity reduced floral displays, including fewer inflorescences and fewer flowers per inflorescence. A reduced floral display was correlated with reduced sucrose, fructose and glucose and resulted in a reduced number and species richness of insect visitors. Fewer flowers per inflorescence reduced the frequency of bumble bee and fly visitors, which were two important pollinators. Although honeybees, flies, small bees, soldier beetles and bumble bees were equally effective pollinators, only bumble bee frequency was positively correlated with pod production. The differences in pollinator visitation have the potential to create diversifying selection on plant floral traits, many of which are also affected by herbivores. This research demonstrates potentially conflicting selection pressures between native and non-native pollinators as well as non-native herbivores.

Melissopalynology: The Louveaux method on honey samples in the marche region and their allergenic potential

The research arises from the interest in investigating the pollen contained in the various honey samples analyzed to identify its geographical and botanical origin and to evaluate its allergenic potential. With the Louveaux method, the pollen content shall be observed under a microscope using the same procedure as the aerobiological analysis described in national and European standards. From the microscopic observation of the pollen grains, it is possible to identify the plants that produced them, thus going back to a particular type of vegetation that distinguishes the production area. The percentage values for pollen of nectariferous plants identified have been calculated to establish frequency classes and dominant species. Starting from an exploratory analysis, the results obtained with melissopalynology provide assessments that, combined with sensory and physical analysis, represent a useful control tool to address the growing trend of food fraud as an element of consumer choice. By comparing the percentages of the pollen taxa contained in the honey analyzed with those reported in the characterization cards, it was possible to verify the conformity to the declared botanical origin. The samples were analyzed in duplicate to reduce operator uncertainty and to ensure the identification of the largest number of species/families present and the recognition of botanical families. The results provided a clear picture of a first assessment of the conformity of the reference values; in general, most of the samples examined, both those taken on the market and those of local producers, are in line with the data obtained. Bibliographical research on cross allergies related to the presence of pollen in honey has established that allergic forms to honey are still a rare condition and the incidence does not exceed 0.001%. Studying the few reported cases worldwide, it is difficult to say that pollen proteins are the only and directly responsible for the allergic manifestations that follow the ingestion of honey; however, if we wanted to attribute the responsibility, we could refer in particular to Compositae pollens, such as mugwort, ragweed, and dandelion. This work, while identifying many possible triggers for allergic reactions, has detected a very low number of cases of ingestion of honey; therefore, the study can be a starting point to investigate the allergenic potential.

Ineffectiveness of ants in protecting two sympatric myrmecophilous plants against endophytic beetles

AbstractExtrafloral nectarized plants attract ants, which may protect them against herbivory and increase plant fruit set production. In some cases, however, ants are ineffective against herbivores. Such events occur, for instance, when herbivores present adaptations to avoid ant predation. Thus, the outputs of these interactions depend on factors such as ant identity, plant phenology and herbivore features. Here, we investigated the endophytic florivorous beetles' impact on the reproductive success of their host plants, two sympatric Banisteriopsis (Malpighiaceae) species, depending on the action of extrafloral nectaries (EFNs) visiting ants. We experimentally manipulated the presence of ants and herbivores on B. malifolia and B. laevifolia species during their reproductive period to test the effectiveness of ants‐EFNs mutualism on plant fruit set production. We performed treatments where three similar inflorescences were selected to be (I) control branch, with no manipulations, (II) beetle‐free branch, isolated with textile cover and (III) ant‐free branch, isolated at the base with atoxic wax. We hypothesized that endophytic herbivores have a negative impact on the plant reproductive success because they escape the action of ants (biotic defences). As a result, we observed (i) that florivorous Anthonomus beetles have a negative impact on Banisteriopsis reproductive success; (ii) ants fail in protecting plants against floral endophytic beetles; and (iii) most of the results were species‐specific. Our results indicate that these systems present very conditional outcomes that depend on the intrinsic factors of each plant species.

A Gis-Based Species Distribution Model for the Endangered Smith's Blue Butterfly, Euphilotes enoptes smithi (Lycaenidae)

A species distribution model (SDM) was developed for the endangered Smith's Blue butterfly, Euphilotes enoptes smithi (Lycaenidae), to predict target areas where it might be found in the rugged Santa Lucia Mountains that extend for 230 km along the Big Sur coast in Monterey County, CA. Field surveys identified 1,506 locations of the butterfly's larval foodplant and primary adult nectar plant, Eriogonum parvifolium (Polygonaceae), in this mountain range. Vegetation, soil, and geology types were identified for each of these known locations. Because all life stages of the endangered butterfly are closely associated with this foodplant, it served as a surrogate to identify new locations that might support the butterfly. Binary logistic regression analysis was used to evaluate the relationships between foodplant occurrence and each of these environmental attributes. Logistic regression identified specific soil and geology types that were useful predictors of potential foodplant occurrence, but it was unable to identify comparable specific vegetation types. GIS analyses were performed to identify target areas that share the same combinations of these environmental attributes in locations known to support the foodplant. The top target areas were all characterized by soil and geology types that the logistic regression analyses corroborated were the best predictors of potential foodplant occurrence. To validate our SDM, field surveys for the foodplant and butterfly were performed at 300 predicted target locations. The foodplant and butterfly were observed at 88.4% and 80.7% of target areas or immediately adjacent to them, respectively. Results of this SDM not only provide a better understanding of this endangered butterfly's potential and actual geographic range but also the environmental attributes that constitute its suitable habitat. Furthermore, these findings will assist resource agencies in their conservation efforts to benefit this endangered butterfly.

Which traits optimize plant benefits? Meta‐analysis on the effect of partner traits on the outcome of an ant–plant protective mutualism

Abstract Theoretical models on mutualism dynamics predict that partner traits may influence the outcome of mutualistic interactions. However, most empirical data on this issue is restricted to case studies, limiting our ability to reach a more widespread comprehension of the role of partner traits on the dynamic of mutualisms. We investigated how the outcome of protective mutualisms between ants and plants bearing extrafloral nectaries (EFNs) is influenced by the traits of EFNs and ants feeding on EFNs. We used a meta‐analytical approach based on 35 studies investigating the effect of ant attendance on the herbivores and reproductive performance of EFN‐bearing plants. We evaluated how variation in the EFN vascularization and location on plants and the ant aggressiveness can modulate the effect of ant attendance on the plants. Both plant and ant traits investigated here drove the outcome of the protective mutualism for EFN‐bearing plants. Plants exclusively bearing EFNs near reproductive organs benefited more from ant attendance than plants bearing EFNs on vegetative or vegetative and reproductive organs. Ants had a higher positive impact on the reproductive performance of plants bearing non‐vascularized EFNs than plants bearing vascularized EFNs, although their effects on herbivores had been similar in both plant types. Regarding the ant behaviour, plants often attended by more aggressive ant species had a higher reproductive performance than plants often attended by less aggressive ones. Synthesis. Our results highlight that the selective pressures and evolutionary routes in ant‐plant protective mutualisms may depend on the pool of traits exhibited by partner species. Although some studies have already reported some impact of species traits on the outcome of ant‐plant mutualisms, this is the first time that a generalization about the role of species traits on the net balance of ant attendance was proposed. Due to this generalization, it was possible to advance our knowledge about the evolution of facultative mutualisms by showing that the role of species traits on the mutualistic outcome can vary in intricate ways due to a particular trait combination found among partners in communities where the interactions are embedded in.

A double defensive mutualism? A case between plants, extrafloral nectaries, and trophobionts

Not applicable

Plants with extrafloral nectaries share indirect defenses and shape the local arboreal ant community.

Associational resistance (AR) is a positive interaction in which a plant suffers less herbivoredamage due to its association with a protective plant. Here, we evaluated whether plants with extra-floral nectaries (EFNs) can share indirect defenses with neighboring plants. We sampled 45 individuals of an EFN-bearing liana (Smilax polyantha) and recorded whether their support species had EFNs. In S. polyantha, we measured foliar herbivory and flower and fruit production. We examined the ant species composition and visitation of S. polyantha and whether they changed according to the supporting plant type (with or without EFNs). We experimentally determined whether S. polyantha supplemented with artificial nectaries could share indirect defenses with defenseless neighboring plants. Support plants with EFNs indirectly benefited S. polyantha by sharing mutualistic ant species. Smilax polyantha supported by plants with EFNs had a more specific ant species composition, a higher number of visiting ants and ant species richness, and exhibitednearly 3 times less foliar herbivory. However, we did not observe differences in fruit production between the two groups of S. polyantha. Finally, we observed that S. polyantha with artificial nectaries increased ant visitation on neighboring plants 2.5 times. We provide evidence that interspecific neighbors with EFNs can experience reciprocal benefits by sharing indirect defenses. Such local effects might escalate and affect the structure of plant communities.

Does the Floral Nectary in Dracocephalum moldavica L. Produce Nectar and Essential Oil? Structure and Histochemistry of the Nectary.

Dracocephalum moldavica is an aromatic plant with a lemon scent and versatile use. Its flowers produce large amounts of nectar, which is collected by bees and bumblebees. The aim of the study was to investigate the structure of the floral nectary in this melliferous plant, which has not been analysed to date. The analyses were carried out with the use of light, fluorescence, scanning electron, and transmission electron microscopy, as well as histochemical techniques. The four-lobed nectary with a diameter of 0.9-1.2 mm and a maximum height of 1.2 mm is located at the ovary base; one of its lobes is larger than the others and bears 20-30 nectarostomata and 8-9 glandular trichomes. The histochemical assays revealed the presence of essential oil and phenolic compounds in the nectary tissues and in glandular trichomes. The nectary tissues are supplied by xylem- and phloem-containing vascular bundles. The nectariferous parenchyma cells have numerous mitochondria, plastids, ribosomes, dictyosomes, ER profiles, vesicles, thin cell walls, and plasmodesmata. Starch grains are present only in the tissues of nectaries in floral buds. The study showed high metabolic activity of D. moldavica nectary glands, i.e., production of not only nectar but also essential oil, which may increase the attractiveness of the flowers to pollinators, inhibit the growth of fungal and bacterial pathogens, and limit pest foraging.

Determination of volatile compound profiles and physico-chemical analysis of linden and acacia Czech honey

In this study 12 acacia and 12 linden honey samples from the Czech Republic were analyzed with a chemometric and volatile compound profiles approach. The results of electrical conductivity (17.4 and 65.1 mS/m), free acidity (12.5 and 24.5 meq/kg), diastase activity (12.6 and 18.8 DN), colour (21.5 and 47.8 mm Pfund), hydroxymethylfurfural (HMF) (11.5 and 3.5 mg/kg), fructose (45.1 and 36.3 g/100g), fructose + glucose (F + G) (76.7 and 69.3 g/100g) content and fructose/glucose ratio (1.4 and 1.1) showed statistically significant differences (p < 0.05) between acacia and linden honey, respectively. The physico-chemical properties and pollen profiles were in compliance with the description of European monofloral honey, except for acacia honey due to the low proportion of acacia pollen (2.8 − 10.3%). According to volatile compounds analysis by head-space solid phase micro-extraction coupled with gas chromatography − mass spectrometry (HS-SPME/GC-MS), nine key volatile substances were selected from more than 100 volatile compounds present in the analysed samples for evaluation as markers of botanical origin typical of the most common plant nectar sources in the Czech Republic. Further, the non-targeted analysis combined with Principal Component Analysis (PCA) revealed trends and outliers within all chromatography data obtained for 24 honey samples. Cis-linalool oxide and β-linalool have been confirmed as markers of acacia honey, while carvacrol and p-cymenene were found almost exclusively in linden honey. The results demonstrated that the chemometrics with the non-targeted and targeted volatile compounds analysis approach is reliable and accurate for the classification of honey of various botanical origins.

Extrafloral nectaries in Nephelium lappaceum (Sapindaceae)

Background: There is no previous evidence in the literature that documents the presence of extrafloral nectaries (EFNs) or the exomorphology of domatia in Nephelium lappaceum (rambutan), a fruit tree native to Malaysia introduced in various American countries.&#x0D; Question: It is the presence of EFNs responsible for the frequent visit of ants to young leaves of Nephelium lappaceum?&#x0D; Study site and dates: This study was conducted in a commercial rambutan orchard in the town of Francisco I. Madero, municipality of Frontera Hidalgo (Chiapas, Mexico) during the period September 2020-February 2021.&#x0D; Methods: External morphology of EFNs and domatia were studied using scanning electron microscopy. Nectar sugar concentration produced by EFNs was determined by refractometry. EFNs location on the plant and the frequency of visiting ants were quantified by visual counting.&#x0D; Results: A description of the external morphology of EFNs and domatia, as well as their location on rambutan leaflets are presented. A volume of 0.3 ± 0.2 µL/EFN of a nectar (9.3 ± 0.6 °Bx) was collected each day. Ten species of ants feeding on the nectar excreted by the EFNs were identified.&#x0D; Conclusion: Evidence of the presence of NEFs on rambutan leaflets is presented, which have an external morphology and location different from that observed in domatia, producing a sweet nectar used as food by a large community of ants.

Extrafloral nectar as entrée and elaiosomes as main course for ant visitors to a fireprone, mediterranean-climate shrub.

Thousands of plants produce both extrafloral nectaries (EFNs) on their leaves and nutrient-rich appendages on their diaspores (elaiosomes). Although their individual ecology is well-known, any possible functional link between these structures has almost always been ignored. Here, we recognized their co-presence in the shrub, Adenanthos cygnorum (Proteaceae), and studied their function and interaction. We observed that the same ants frequently visit both structures, seeds are attractive to vertebrate granivores but are released into a leafy cup from where they are harvested by ants and taken to their nests, from which seeds, lacking elaiosomes, germinate after fire. We showed that juvenile plants do not produce EFNs and are not visited by ants. We conclude that EFNs are not just an indirect adaptation to minimize herbivory via aggressive ant visitors (the role of a minority) but specifically enhance reproductive success in two ways: First, by inducing ants to visit the plant as a reliable food source throughout the year. Second, by promoting discovery of the seasonally available, elaiosome-bearing seeds for transport to their nests (the majority of visitors), so avoiding the risk of granivory should seeds instead fall to the ground. Parasitoid wasps play a supporting role in controlling the main insect herbivore whose larvae devour the reproductive apices. Thus, the EFN-elaiosome relationship has three components that enhance species fitness: foliage protection, seed transport, and granivore escape. A similar system has been described only once before (in an unrelated biome) and, consistent with the objectives of ecology as an integrative science, deserves wider study.

Extrafloral nectaries in Passiflora pohlii Mast. (Passifloraceae): morpho-anatomical and ecological aspects of an understudied nectary morphotype in Passiflora L.

Extrafloral nectaries in Passiflora pohlii Mast. (Passifloraceae): morpho-anatomical and ecological aspects of an understudied nectary morphotype in Passiflora L.

Precision Phenotyping of Nectar-Related Traits Using X-ray Micro Computed Tomography.

Flower morphologies shape the accessibility to nectar and pollen, two major traits that determine plant-pollinator interactions and reproductive success. Melon is an economically important crop whose reproduction is completely pollinator-dependent and, as such, is a valuable model for studying crop-ecological functions. High-resolution imaging techniques, such as micro-computed tomography (micro-CT), have recently become popular for phenotyping in plant science. Here, we implemented micro-CT to study floral morphology and honey bees in the context of nectar-related traits without a sample preparation to improve the phenotyping precision and quality. We generated high-quality 3D models of melon male and female flowers and compared the geometric measures. Micro-CT allowed for a relatively easy and rapid generation of 3D volumetric data on nectar, nectary, flower, and honey bee body sizes. A comparative analysis of male and female flowers showed a strong positive correlation between the nectar gland volume and the volume of the secreted nectar. We modeled the nectar level inside the flower and reconstructed a 3D model of the accessibility by honey bees. By combining data on flower morphology, the honey bee size and nectar volume, this protocol can be used to assess the flower accessibility to pollinators in a high resolution, and can readily carry out genotypes comparative analysis to identify nectar-pollination-related traits.

Croton (Euphorbiaceae) of the Brazilian state of Paraná: an annotated checklist, species distribution, and identification key

Croton is a megadiverse pantropical genus with about 1200 species. The genus can be easily recognized by a set of characters that include indumentum of stellate or lepidote trichomes, colored or clear latex, leaves often with nectary glands, and terminal inflorescences. We focus on the State of Paraná in southern Brazil, where we identified a gap in knowledge of Croton. This study was based on fieldwork and herbarium surveys that allowed us to document all Croton species occurring in the state. We recognize 41 species for the state and provide an identification key, taxonomic notes, and distribution maps for all species. We also include photographs and carry out lectotypifications when necessary. We expect this work will contribute to a better understanding of Croton diversity and distribution in southern Brazil.

THE FIRST EXPERIENCE IN THE DEVELOPMENT OF THE RICHES OF THE VASYUGAN SWAMP

Introduction to Siberia of bees of different breeds such as Italian Apis mellifera ligustica Spin., Apis mellifera carnica Pollm., Kuban (yellow Caucasian) Apis mellifera remipes Gerst., Far Eastern Apis mellifera acervorum, grey mountain Caucasian Apis mellifera caucasica Gorb. led to the fact that crossbreeds of an unknown generation predominate in the Novosibirsk region. These bees are rich in swarms, do not winter well, and the egg-laying of the queens is low. Currently, the importation of packet bees has stopped due to sharp increases in fuel prices (fuels and lubricants) and air and rail transport. But the importation of fetal queen bees is still ongoing. The authors studied the method of reproduction of bee colonies to preserve the beekeeping industry in Western Siberia. In one of the promising regions of Western Siberia, in the Vasyugan swamps, nectariferous plants grow for breeding bees. In that place, plants grow such as Rubus idaeus L. (common raspberry), Angelica sylvestris L. (wood angelica), Archangelica decurrens L. (Siberian angelica), Ribus nigrum L. (black currant), Sonchus arvensis L. (field sow thistle), Melilotus officinalis L. (yellow sweet clover), Melilotus albus Medik. (White sweet clover), Centaurea scabiosa L. (rough cornflower), Salix L. (willow), Taraxacum Wigg. (dandelion), Dracocephalum nutans L. (Siberian snakehead), Glechoma bederaceae L. (ivy-shaped budra), Prunella vulgaris L. (common blackhead). These plants are an excellent forage base for beekeeping. For two years of research, the authors found that the collection of honey per bee colony was in the group where the bees were propagated by natural swarming, 51.8 ± 5.1 kg, individual layering - 49.9 ± 4.3, prefabricated layering - 55, 6±3.1 kg. The strength of the families that went into the winter was 8.47±0.30; 8.77±0.20 and 9.06±0.40 bee space, respectively. Reproduction of bee colonies for two years showed that it is possible to increase the number of bee colonies from 4 to 16 (403%) by natural swarming, by individual layers - up to 19 (572%), and teams - up to 17 (422%).

Yearly weather variation and surface temperature drives the spatiotemporal dynamics of a threatened butterfly and its host plant

It remains unclear to what extent yearly weather variation and spatial variation in microclimate influences the outcome of interacting plant-animal species and whether responses differ between life stages. We collected data over several years on 46 ha on File Hajdar, Gotland, Sweden, and executed a complete mapping of larva nests (n= 776) and imago (n= 5,952) of the marsh fritillary butterflyEuphydryas auriniaand its host plantSuccisa pratensis. The phenology of the butterflies and the major nectar plants visited varied among years. The duration of the adult flight period decreased with increasing ambient air temperatures. The density of butterflies, host plants, and host plant leaf size increased between years with increasing precipitation in the preceding year, and decreased with increasing average ambient air temperature in the preceding year. In 2021–2022 we deployed a unmanned aerial vehicle (UAV) with a high-resolution thermal sensor to measure spatial variation in surface temperatures in the study area. We found that survival from the egg to the larva stage increased with increasing surface temperature and host plant density. Host plants and larva nests generally occupied warmer microhabitats compared to imago butterflies. The results further suggested that the relationships linking surface temperature to the densities of imago, larva, host plants, and leaf size differed qualitatively between years. In 2017, larva nests and host plant density increased with increasing surface temperatures, and butterflies showed a non-linear response with a density peak at intermediate temperatures. As a result of the extreme drought in 2018 there was a reduction in maximum leaf size, and in the densities of plants, larvae, and butterflies. Moreover, the slopes of the relationships linking the density of larvae, butterflies, and plants to temperature shifted from linear positive to negative or curvilinear. Our findings demonstrate how yearly weather variation and heterogeneous surface temperatures can drive the spatiotemporal distribution and dynamics of butterflies and their host plants. The context specificity of the responses indicated by our results makes it challenging to project how climate change will affect the dynamics of ecological communities.

Touch-sensitive stamens enhance pollen dispersal by scaring away visitors.

Animal-pollinated plants have to get pollen to a conspecific stigma while protecting it from getting eaten. Touch-sensitive stamens, which are found in hundreds of flowering plants, are thought to function in enhancing pollen export and reducing its loss, but experimental tests are scarce. Stamens of Berberis and Mahonia are inserted between paired nectar glands and when touched by an insect's tongue rapidly snap forward so that their valvate anthers press pollen on the insect's tongue or face. We immobilized the stamens in otherwise unmodified flowers and studied pollen transfer in the field and under enclosed conditions. On flowers with immobilized stamens, the most common bee visitor stayed up to 3.6× longer, yet removed 1.3× fewer pollen grains and deposited 2.1× fewer grains on stigmas per visit. Self-pollen from a single stamen hitting the stigma amounted to 6% of the grains received from single bee visits. Bees discarded pollen passively placed on their bodies, likely because of its berberine content; nectar has no berberine. Syrphid flies fed on both nectar and pollen, taking more when stamens were immobilized. Pollen-tracking experiments in two Berberis species showed that mobile-stamen-flowers donate pollen to many more recipients. These results demonstrate another mechanism by which plants simultaneously meter out their pollen and reduce pollen theft.

Genetic architecture of multiple mutualisms and mating system in Turnera ulmifolia.

Plants often associate with multiple arthropod mutualists. These partners provide important services to their hosts, but multiple interactions can constrain a plant's ability to respond to complex, multivariate selection. Here, we quantified patterns of genetic variance and covariance among rewards for pollination, biotic defence and seed dispersal mutualisms in multiple populations of Turnera ulmifolia to better understand how the genetic architecture of multiple mutualisms might influence their evolution. We phenotyped plants cultivated from 17 Jamaican populations for several mutualism and mating system-related traits. We then fit genetic variance-covariance (G) matrices for the island metapopulation and the five largest individual populations. At the metapopulation level, we observed significant positive genetic correlations among stigma-anther separation, floral nectar production and extrafloral nectar production. These correlations have the potential to significantly constrain or facilitate the evolution of multiple mutualisms in T.ulmifolia and suggest that pollination, seed dispersal and defence mutualisms do not evolve independently. In particular, we found that positive genetic correlations between floral and extrafloral nectar production may help explain their stable coexistence in the face of physiological trade-offs and negative interactions between pollinators and ant bodyguards. Locally, we found only small differences in G among our T.ulmifolia populations, suggesting that geographic variation in G may not shape the evolution of multiple mutualisms.