Presence Of Extrafloral Nectaries Research Articles

Young stem and leaf anatomy of 15 Malagasy-endemic Diospyros species (Ebenaceae): taxonomic implications

Abstract Eighty-eight of the 255 Diospyros species from Madagascar are potential sources of ebony wood. Unfortunately, several species are currently threatened with extinction due in part to illegal and unsustainable logging. Reliable identification of living Diospyros species is necessary prior to any sustainable exploitation, so as not to compromise those species threatened with extinction. This study aims to identify features that might contribute to supporting taxonomic studies and to provide a complementary tool for the identification of standing trees. Fifteen species of Diospyros endemic to Madagascar were studied using standard anatomical methods. Each species showed a different shape of the leaf midrib and some have distinctive leaf anatomical characteristics, such as the presence of extrafloral nectaries in the distal part of the petiole in Diospyros baronii and filiform sclerenchyma in the mesophyll in Diospyros lewisiae. Statistical analysis showed that petiole and midrib vascular patterns, spongy parenchyma structure, the presence of papillae, thickness of epidermis, presence of extrafloral nectaries, stratification of epidermis, and presence of a cuticle are useful to distinguish species groups and even some species. The anatomy of young stems and leaves of Diospyros species from Madagascar is examined for the first time; the study will be extended to other potentially exploitable species.

Experimental manipulation of biotic and abiotic parameters changes the outcome of insect-plant interactions

Plants usually respond to environmental shifts with morpho-physiological adjustments, which trickles down to biotic interactions in the insect-plant system. We evaluate how Copaifera langsdorffii, a widespread tree species adapted to multiple ecosystems, responds to shifts in nutrient and water availability through experimental supplementation and how it affects its insect communities. We also evaluate how the presence of extrafloral nectaries (EFN) exudates modifies galling insect diversity and herbivory rates. Such experimental approach is scarce, as we simultaneously evaluate biotic and abiotic factors and cover the entire reproductive cycle of a species in natural environments, bringing important contributions to better understand how bottom-up factors drive species interactions and possibly community assembly. The experiment was set in an ironstone outcrop vegetation, where we deployed supplementation treatments in 80 plant individuals divided into the following field treatments: (T1 = Fertilizer, T2 = EFN simulation (Eppendorf tubes with a 20% sugar solution), T3 = fertilizer + EFN simulation, T4 = water spray, T5 = EFN control (microtube with water), and T6 = Control. We observed lower sclerophylly and greater leaf area in plant individuals that were supplemented with nutrients and water. Herbivory rates were lower and ant abundance was higher in plants with artificial EFNs available. While we did not observe variations in richness and abundance of galling insects, the patterns of co-occurrence of galls varied with the availability of resources (nesting space) in the plant. This study illustrates how variations in nutrient availability to plants modify interactions with insect communities. Ant-plant interactions can have a negative impact on general herbivory rates, however ants seem to have a more harmonious relationship with the galling insects.

Alien and native tree species having extrafloral nectaries as favorite hunting area for arboreal endemic Philippine tiger beetles (Coleoptera: Cicindelidae) in human-disturbed habitat in Lanao del Sur Province, Mindanao, Philippines.

To document a relation between abundance of arboreal, predatory tiger beetles, their ant prey, and extrafloral nectaries attracting the ants, we gathered data from more than 10 species of native and introduced trees and large, tree‐like perennial plants in Lanao del Sur Province, Mindanao, Philippines. All specimens of tiger beetles (two Tricondyla and two Neocollyris species, all endemic to the country) were noted on five tree species characterized by presence of extrafloral nectaries, including three alien/invasive and two native ones. Invasive Spathodea campanulata and native Hibiscus tiliaceus were the most inhabited ones (respectively, 56% and 19% of beetles). Presence of tiger beetles on these trees most probably depends on high abundance of ants, which are typical prey for arboreal Cicindelidae, while occurrence of ants can result from presence of extrafloral nectaries on different parts of the plants. This suggests a new mutualistic insect–plant interaction between native and invasive species.

Ant species richness and interactions in canopies of two distinct successional stages in a tropical dry forest.

Canopy ecology is a fast-growing field, but still a scientific frontier in many ecological aspects. For instance, the hypothesis that tree traits shape patterns in ant-plant interactions lacks data, notably for tropical canopies in different successional stages. In this study, we investigated canopy traits, such as tree height, the presence of extrafloral nectaries (EFNs), connectivity among tree crowns, and successional stage, structure ant-tree interactions in a tropical dry forest (TDF), examining whether these are the determinant factors for ant species richness. We collected ants on trees in early and late successional stages over 2years, in rainy and dry seasons. In the late successional stage, ant species richness was greater in the taller trees; in the early successional stage, the smallest trees had a greater ant species richness than the taller trees. The EFNs and connectivity among treetops had no effect on ant species richness. We obtained a tree-ant network of the early successional stage, involving 786 interactions among 57 ant species and 75 trees; in the late successional stage, the network had 914 interactions among 60 ant species and 75 trees. There were 27 species of trees in our study, 11 of which (40.7% of all individual trees) had EFNs. The ant-plant interactions were not randomly distributed, suggesting that various biotic factors structured the ant assemblies. This study presents new insights into ant-tree interactions, showing that both tree height and successional stage influence the occurrence of many species of ants in tree canopies of tropical dry forests.

Morpho–anatomical and ultrastructural analysis of extrafloral nectaries in Inga edulis (Vell.) Mart. (Leguminosae)

The presence of extrafloral nectaries (EFNs) between leaflets is an usual feature in Inga edulis (Vell.) Mart. (Leguminosae). Extrafloral nectaries are secretory structures involved in production of nectar and which serve in the protection of plants against herbivores through association with ants. This study aimed to characterize the EFNs of I. edulis at different developmental stages and describe their morphology, histochemistry and ultrastructure. Leaf fragments, containing secretory structures, were processed according to standard methods for light, scanning and transmission electron microscopy. The EFNs were classified into three stages based on morphology: pre‐secretory, secretory and post‐secretory. The pre‐secretory stage occurs in young leaves, whereas secretory and post‐secretory stages occur in developed and senescent leaves, respectively. The EFNs possess a concave surface and a central cleft in which nectar is accumulated and which was not observed in pre‐secretory EFNs. Histochemical tests identified the presence of sugars, proteins, phenolic compounds, mucilage and lipids at all developmental stages of the EFNs. Calcium crystals were identified in all tissues and stages of the EFNs. The secretory cells of the EFNs exhibit a granular cytoplasm, small vacuoles, prominent nuclei, smooth endoplasmic reticulum and mitochondria. Post‐secretory stage EFNs exhibited intense cytoplasmic degradation and the presence of microorganisms. The performance of EFNs of I. edulis appear to follow the leaf development.

Extrafloral nectar as a driver of arboreal ant communities at the site‐scale in Brazilian savanna

AbstractDespite years of study, it remains unclear if and to what extent the effects of extra‐floral nectaries (EFNs) on arboreal ants observed on individual trees scale up to larger spatial scales. Here, we address this issue in Brazilian savanna and tested three predictions: (i) Trees with EFN have higher richness of arboreal ant species than trees without; (ii) Arboreal ant species richness increases with the proportion of total EFN‐bearing trees at the site scale, due to a higher occurrence of non‐core ant species; (iii) Ant species composition changes with the proportion of EFN‐bearing trees at the site scale. We sampled arboreal ants in 32 plots with EFN‐bearing trees ranging from 0% to 60% of all trees. We sampled 72 ant species, from which 17 (mostly belonging to Camponotus, Cephalotes and Crematogaster) were identified as core species in at least one of the ant‐EFN networks in the 32 plots. Ant species richness was significantly higher on EFN‐bearing trees. We identified 11 ant species that preferentially occurred on EFN‐bearing trees, all of which were core partners in networks. Species richness at the site scales increased with the proportion of EFN‐bearing trees, regardless of tree density and richness; this pattern was due to a higher occurrence of non‐core ant species. Finally, species composition also varied with the proportion of EFN‐bearing trees. Therefore, we found that the presence of EFNs not only influences arboreal ants on individual trees but also has a substantial effect on the ant‐EFN network on a broader community scale. The increase in non‐core species site scale reveals that this interaction is unlikely to result in substantially enhanced protection services for EFN‐bearing plants.

Leguminosae clado mimosoide em um fragmento de floresta estacional semidecidual do sudeste do Brasil

Resumo A floresta estacional semidecidual, fitofisionomia que revestia quase todo o interior do estado de São Paulo, está representada atualmente por apenas 5% da sua cobertura original. Este trabalho consiste no levantamento dos representantes do clado mimosoide numa das últimas áreas de vegetação contínua do estado, a Serra do Japi. Foram registradas 24 espécies distribuídas em oito gêneros, dos quais os mais diversos são Mimosa (7 spp.), Inga (6 spp.), Senegalia (5 spp.) e Anadenanthera (2 spp.). As espécies de hábito arbóreo predominam no local e a maioria dos táxons compartilha o Cerrado e a Mata Atlântica como área de ocorrência, com exceção de Inga subnuda e Senegalia grandistipula, ambas endêmicas da Mata Atlântica. Seis são endêmicas do Brasil (Abarema langsdorffii, I. subnuda., I. sessilis, Leucochloron incuriale, Mimosa furfuracea, S. grandistipula), doze são novos registros para a Serra do Japi e uma é inédita para o estado (Senegalia tucumanensis). Os principais caracteres diagnósticos para a distinção dos gêneros foram os tipos de folha e fruto, presença de nectários extraflorais e número e fusão de estames, e para as espécies, presença de glândulas em locais específicos, presença de acúleos, tricomas e estípulas, e número de folíolos.

Sugar secretion and ant protection in Ficus benguetensis: Toward a general trend of fig–ant interactions

The relationship between plants and ants is often mediated by the presence of extrafloral nectaries (EFNs) that attract ants and provide rewards by protecting plants from herbivores or parasites. Ficus trees (Moraceae) and their pollinators (Hymenoptera: Agaonidae) are parasitized by many nonpollinating fig wasp species (Hymenoptera: Chalcidoidea) that decrease the reproductive output of the mutualistic partners. Previous studies have shown that ants living on and patrolling Ficus species can efficiently deter parasitic wasps. The aim of this study was to verify the presence of EFNs on figs of Ficus benguetensis and test the hypothetical protection service provided by ants. Figs in different developmental stages were collected from Fu-Yang Eco Park, Taipei, Taiwan. Sugars on the fig surface were collected and analyzed through high-performance anion-exchange chromatography. Moreover, ants were excluded from the figs to determine the effect of ants on the nonpollinating fig wasps. We identified three oligosaccharides whose relative proportions varied with the fig developmental phase. In addition, results showed that the ant-excluded figs were heavily parasitized and produced three times less pollinators than did the control figs. Finally, the specific interactions of Ficus benguetensis with ants and the relationship between figs and ants in general are discussed.

Ants visiting extrafloral nectaries and pyrrolizidine alkaloids may shape how a specialist herbivore feeds on its host plants

The presence of extrafloral nectaries (EFNs) attracts predators and parasitoids, and protects the plant against herbivorous insects. By improving plant defences, EFNs reduce the fitness of herbivores. The use of similar host plants with no EFNs or adaptations in response to predators and parasitoids may enhance herbivore fitness. In this context, we studied the feeding habit (on leaves or on unripe seeds inside the pods) of larvae of the specialist moth Utetheisa ornatrix in two Crotalaria host plant species in which EFNs are present (C. micans) or absent (C. paulina). We hypothesized that the moths’ feeding habit was influenced by its natural enemies via their presence on EFNs. In C. micans, we found more larvae feeding inside the pods rather than on the leaves, while in C. paulina, larvae were found in both parts of the plant. There was greater activity of natural enemies in C. micans than in C. paulina. The moth sequesters enough pyrrolizidine alkaloid (PAs) to defend against predators in the leaves and seeds of C. paulina, but only in seeds of C. micans. Therefore, a change in the feeding habit in U. ornatrix larvae is a plastic response that depends on whether EFNs are present or not, or whether PA concentrations are low or high. This change does not affect overall moth performance. However, other factors, such as pod hardness, predation by organisms other than those visiting EFNs or even parasitoids cannot be ruled out as being responsible for the change in feeding habit. To date, both the EFNs and PAs in Crotalaria species are a parsimonious explanation of how larvae of U. ornatrix use different species of Crotalaria for feeding.

Decreasing water availability across the globe improves the effectiveness of protective ant-plant mutualisms: a meta-analysis.

Abiotic conditions can increase the costs of services and/or the benefits of rewards provided by mutualistic partners. Consequently, in some situations, the outcome of mutualisms can move from beneficial to detrimental for at least one partner. In the case of protective mutualisms between ant bodyguards and plants bearing extrafloral nectaries (EFNs), plants from arid environments face a trade-off between EFN production and maintenance and water and carbon economy. This trade-off may increase EFN costs and decrease their value as a defensive strategy to plants in such environments. Despite this, the presence of EFNs is an ubiquitous trait in plants from arid environments, suggesting that they provide greater benefits to plants in these environments to compensate for their higher costs. We used a meta-analysis to investigate if such benefits do increase with decreasing water availability and the possible underlying causes (such as ant behaviour or ant diversity). As predicted, ant effect on EFN plants performance increased as mean annual precipitation decreased. We also found that the frequency of dominant ants on EFN plants increased in drier areas. Due to the more aggressive behaviour of dominant ants, we suggest that they represent an important factor shaping the adaptive value of EFNs to plants in arid environments.

Extrafloral nectaries inPhilodendron(Araceae): distribution and structure

Extrafloral nectaries (EFNs) are involved in animal–plant interactions that lead to protection against herbivory. The presence of EFNs in Araceae is rare, besides Philodendron, there is report for only two other genera. With the aim to investigate the occurrence of EFNs in Philodendron and to describe the distribution patterns and structural organization of these glands, 75 Philodendron spp. were examined, 16 of which were selected for study by light microscopy. Three Homalomena spp. were also examined for EFNs, but these were not found. Philodendron martianum was employed as a model for additional study using scanning and transmission electron microscopy. The studied EFNs showed a high degree of structural similarity. They were present in the prophyll, leaf and spathe, becoming functional in young organs. In surface view, EFNs consisted of small areas and showed one or more stomata through which secretions were released. The secretory cells formed a globular region surrounded by ground parenchyma. In P. martianum, nectariferous parenchyma cells exhibited typical features of cells with high metabolism related to nectar secretion. These results allow us to infer that EFNs have a widespread occurrence in Philodendron, and they remain an exclusive character for this group. © 2015 The Linnean Society of London, Botanical Journal of the Linnean Society, 2015, ●●, ●●–●●.

How territoriality and host-tree taxa determine the structure of ant mosaics.

Very large colonies of territorially dominant arboreal ants (TDAAs), whose territories are distributed in a mosaic pattern in the canopies of many tropical rainforests and tree crop plantations, have a generally positive impact on their host trees. We studied the canopy of an old Gabonese rainforest (ca 4.25 ha sampled, corresponding to 206 "large" trees) at a stage just preceding forest maturity (the Caesalpinioideae dominated; the Burseraceae were abundant). The tree crowns sheltered colonies from 13 TDAAs plus a co-dominant species out of the 25 ant species recorded. By mapping the TDAAs' territories and using a null model co-occurrence analysis, we confirmed the existence of an ant mosaic. Thanks to a large sampling set and the use of the self-organizing map algorithm (SOM), we show that the distribution of the trees influences the structure of the ant mosaic, suggesting that each tree taxon attracts certain TDAA species rather than others. The SOM also improved our knowledge of the TDAAs' ecological niches, showing that these ant species are ecologically distinct from each other based on their relationships with their supporting trees. Therefore, TDAAs should not systematically be placed in the same functional group even when they belong to the same genus. We conclude by reiterating that, in addition to the role played by TDAAs' territorial competition, host trees contribute to structuring ant mosaics through multiple factors, including host-plant selection by TDAAs, the age of the trees, the presence of extrafloral nectaries, and the taxa of the associated hemipterans.

Biological control of the greenhouse whitefly by Amitus fuscipennis: Understanding the role of extrafloral nectaries from crop and non-crop vegetation

To determine the potential role of non-crop vegetation extrafloral nectaries (EFNs) in the presence and feeding of Amitus fuscipennis, a parasitoid of Trialeurodes vaporariorum, non-crop vegetation in and around ten fields of common beans (Phaseolus vulgaris) was studied in the Central Andes of Colombia. Using 0.25m2 quadrants located randomly both within the bean fields and at a distance of 3m from the crop, the non-crop vegetation was identified and counted. The abundance of both T. vaporariorum and A. fuscipennis was also determined. Furthermore, plants separated by more than 3m from the crop were examined for 30min to determine the presence of A. fuscipennis. There were 92 plant species associated with bean crops: 46 within the crop, 73 at 3m from the crop border, and an additional 18 species at more than 3m from the crop. The occurrence of A. fuscipennis was influenced by the presence of EFNs and whitefly nymphs. Some plants allowed reproduction of both insect species. Experimental data confirmed that A. fuscipennis fed on EFNs. The role of the crop itself as a sugar source for the parasitoid was studied. It was confirmed that P. vulgaris stipels are EFNs and a source of sugar for A. fuscipennis. Several non-crop plant species in the agroecosystem and within the bean crop play a role in the whitefly/parasitoid interaction as alternate hosts for T. vaporariorum and as a source of concentrated sugar in EFNs for A. fuscipennis.

Why eat extrafloral nectar? Understanding food selection by Coleomegilla maculata (Coleoptera: Coccinellidae)

Methods of increasing predator abundance within a habitat include the incorporation of non-prey food items, yet the influence of this on predation intensity toward herbivores remains unknown. In order to gain an understanding of nectar feeding in the predaceous beetle, Coleomegilla maculata (DeGeer), laboratory studies were conducted evaluating prey consumption in the presence of extrafloral nectaries. The physiology of beetles with access to prey only and a mixed diet were compared. To elucidate results of beetle physiology, Y-tube olfactometer studies were conducted and preferences between food types evaluated. Coleomegilla maculata females consumed 9 % fewer aphids when nectar was available. Lipid and glycogen content, as well as oocyte volume were not increased upon consumption of a mixed diet. Evaluation of predator behavior when offered both food resources together and separately demonstrated that extrafloral nectaries are attractive.

Shelter-building caterpillars in the cerrado: seasonal variation in relative abundance, parasitism, and the influence of extra-floral nectaries

Caterpillar shelters provide protection against desiccation and natural enemies, whereas extra-floral nectaries (EFNs) may be an anti-herbivore adaptation that reduces herbivore abundance by attracting predators and parasites. We used a large, long-term dataset for caterpillars found in the Brazilian cerrado to examine temporal variations in the relative abundance of shelter-building caterpillars and exposed caterpillars, and to determine how much variation depends on the season and the presence of EFNs on host plants. We also compared the patterns of parasitism between sheltered and exposed caterpillars, between seasons, and between different host plants. The cerrado has a marked dry season, and its vegetation is a mixture of mostly deciduous shrubs and trees. Leaf production occurs mainly during the rainy season, and many plant species bear EFNs. Our results show that 60 % of cerrado caterpillars build shelters. These caterpillars were found to be proportionally more abundant during the dry season and less parasitized than exposed ones. The proportion of caterpillars building shelters was highest on plants with new leaves (functional EFNs), and parasitism of caterpillars on these plants was higher. Even though our study includes a taxonomically diverse suite of caterpillars that build many different types of shelter and a diverse set of plants and EFN types, our results suggest that EFNs play an important role in structuring caterpillar assemblages in the cerrado, and that the prolific use of shelters by caterpillars may be a result of their effectiveness in protecting caterpillars from natural enemies and desiccation.

Seed fate in the myrmecochorous Neotropical plant Turnera ulmifolia L., from plant to germination

Myrmecochory (seed dispersal by ants) differs from other dispersal systems in a series of advantages offered by the ants to the plants. Here, seed fate, from fruit to germination, of the myrmecochorous Neotropical plant Turnera ulmifolia L. is described. Seed movement from the fruit to their germination was studied, using different measurements and experiments. The results show that a T. ulmifolia individual produces ca. 5000 seeds per year. The main pre-seed-fall predators are the larvae of the Microlepidopteran Crocidosema plebejana Zeller, which consumed 1% of the seeds on the plant. The red-land crab Gecarcinus lateralis (Freminville) consumed 19% of the seeds beneath the plant and was the main post-seed-fall predator. Seed removal by ants was recorded on and beneath the plant, and ants removed 49% of the total seed production. Considering the seed removal events, the ant Forelius analis contributed with 64% of the total number of events. F. analis took seeds to its nest and discarded 23% of the seeds collected. Germination of seeds collected by F. analis was two to four times higher than that of seeds with and without elaiosome, respectively. The relatively low seed predation was probably related to ant defense, associated with the presence of extrafloral nectaries in this plant and with seed removal on the plant. Our results suggest that F. analis is a quantitatively efficient but qualitatively inefficient seed disperser of T. ulmifolia.

Within‐plant distribution of phenolic glycosides and extrafloral nectaries in trembling aspen (Populus tremuloides; Salicaceae)

Expression of foliar secondary compounds and extrafloral nectaries (EFNs) within the same leaves may be incompatible if secondary compounds repel beneficial insects that might otherwise be attracted to EFNs. This study examined the within-plant distributions of phenolic glycosides and EFNs in trembling aspen, Populus tremuloides, and their relationships to herbivore damage. Populus tremuloides expresses extrafloral nectaries (EFNs) on a subset of its leaves. We studied short and tall naturally occurring aspen ramets across multiple sites in interior Alaska. Contrary to our expectations, foliar phenolic glycoside concentrations were approximately 30% greater on leaves bearing EFNs than on leaves without EFNs. The mean concentration of foliar phenolic glycosides in short ramets was nine times that in tall ramets. Phenolic glycoside concentration was negatively related to leaf mining damage by Phyllocnistis populiella (Lepidoptera; Gracilliadae) at concentrations greater than 27 mg/g, whereas the presence of EFNs was unrelated to mining damage. The positive association of chemical defensive compounds and EFNs in leaves suggests that, for species with variation in EFN expression, negative correlations between herbivory and EFN expression may arise indirectly from associated effects of other, correlated types of defense.

The arboreal ant community visiting extrafloral nectaries in the Neotropical cerrado savanna

AbstractThe cerrado savanna of Brazil embraces an area of approximately 2 million km2, in which vegetation physiognomies may vary from open grassland to forest with a discontinuous herbaceous layer. Here we describe the main ecological factors accounting for the prevalence of ants on cerrado foliage, and present a general characterization of the arboreal ant fauna of this savanna. The high incidence of ants on cerrado foliage results mostly from the wide occurrence of predictable liquid food sources in the form of extrafloral nectaries (EFNs) and insect honeydew, which act as efficient promoters of ant activity on vegetation. In addition, stem galleries and cavities constructed by boring beetles and insect galls create a nesting space frequently used by arboreal ants. Specific studies involving ants, herbivores and plants are reported to demonstrate the impact that foliage-dwelling ants can have on phytophagous insects, herbivory levels, and ultimately on host plants. These studies show that: (i) ants visit EFNs and likely benefit from this resource; (ii) EFN-gathering ants can benefit particular plant species by reducing herbivory and increasing plant fitness; (iii) presence of EFNs does not affect ant species richness within a given tree; (iv) there is not a particular ant species composition typical of plants with EFNs; (v) although plants with EFNs are visited by more ant individuals than non-nectariferous plants, this visitation pattern does not translate into lower numbers of herbivores on the nectariferous plant community. We suggest some promising research avenues to elucidate how community-level parameters can be tied to the ecology of ant-plant associations in cerrado.

Effects of grazing and ant/beetle interaction on seed production in the legume Vicia sepium in a seminatural grassland

Abstract 1. Seed production can be affected by cattle grazing, insect herbivores, and seed predators. Bush vetch, Vicia sepium L. (Fabaceae), possess extra floral nectaries (EFNs), which can attract ants that act as plant defenders.2. In this field study, four different manipulations were used to investigate interactions between V. sepium, ants, aphids, and seed predators, mainly Bruchus atomarius. The experiment was set up in grazed and non‐grazed plots in a semi‐natural grassland in south‐central Sweden.3. Grazing negatively affected number of V. sepium buds, flowers, and pods. Seed set, defined as the number of developed seeds per number of ovules in a pod, was higher in grazed plots. Ant abundance, however, was not affected by grazing. Ten different ant species were found on V. sepium and the number of ants was positively correlated with the presence of EFNs. Aphids were found only when ants were excluded from the plants. Pod and seed production of V. sepium plants was not affected by the presence of ants. The seed predator B. atomarius was not affected by the presence of ants. No support was found for the supposition that V. sepium benefited from attracting ants by producing extra floral nectar.4. A small observational study of V. sepium plants in shrub (Rosa dumalis) plots compared with grassland plots was added to the experimental study. Vicia sepium plants were longer and had higher seed‐set (seeds/ovules) in shrub plots than in grassland plots.

Effects of ant behaviour and presence of extrafloral nectaries on seed dispersal of the Neotropical myrmecochore Turnera ulmifolia L. (Turneraceae)

This paper describes the myrmecochory system of Turnera ulmifolia in a coastal sand dune matorral in Mexico. Turnera ulmifolia has elaiosome-bearing seeds and extrafloral nectaries (EFNs). In ten quadrants (4 × 15 m) ant–seed interaction was monitored, and an interaction intensity index calculated and correlated with the number of seedlings. Seed removal rates by ants were surveyed every 2 h for 24 h, the ants being observed both on and beneath the plants. The role of the elaiosome in seed removal was evaluated by offering seeds with and without elaiosomes, and elaiosomes only. Finally, the effect of ant manipulation in seed germination was evaluated. There were 25 ant species associated with seeds and/or EFNs, the most frequently recorded being Monomorium cyaneum and Forelius analis. There was a positive correlation between the intensity index and seedling number per quadrant. There was significantly higher mean seed removal during the day than during the night (19.3% and 12.3%, respectively), and from beneath than on the plant (21.9% and 9.5%, respectively). The preference for elaiosomes only was also greater during the diurnal period, and when gathered on, rather than beneath, the plant. Seed manipulation by F. analis enhanced germination by T. ulmifolia. Seed removal, dispersal distances, seed predation and germination were largely determined by ant behaviour. The presence of EFNs may be influencing seed removal on the plant by attracting a specific assemblage of omnivorous ants. Among such assemblages associated with T. ulmifolia we encountered a variety of behaviours, with ant species either good at defending plants but bad at dispersing seeds, or vice versa. We discuss the way in which these two rewards, and the processes involved (defence and dispersion), could have interacted with each other and evolved. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 86, 67–77.