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.     

Flower nectar of an autogamous perennialRorippa indica as an indirect defense mechanism against herbivorous insects

This report shows that one of the most important roles of the flower nectar of an autogamous perennialRorippa indica (L.) Hieron is as an attractant for employing some ant species as a defense against herbivorous insects. The plant has flowers from spring to early winter. Its flower nectar is frequently stolen by some ant species (hereafter cited as ants) which also feed on small herbivorous insects on the plant. Internations among the tritrophic levels (R. indica, herbivores, ants) were experimentally examined and the followings became clear. (1) Ants were attracted toR. indica in search of its flower nectar. (2) The gradual secretion of flower nectar seemed to detain ants on the plant. (3)Pieris butterfly lavae were the major herbivores onR. indica and were potentially harmful to the plant. (4) The presence of ants reduced the survival rate ofP. rapae larvae onR. indica. (5) The presence of ants reduced the feeding damage toR. indica. (6) The disadvantage of nectar use by ants seemed to be minimal for the plant since the ants did not disturb the other flower visitors. These facts suggest a mutualistic relationship betweenR. indica and ants. That is, the flower nectar serves as an indirect defense against herbivorous insects.     

Various Chemical Strategies to Deceive Ants in Three Arhopala Species (Lepidoptera: Lycaenidae) Exploiting Macaranga Myrmecophytes

Macaranga myrmecophytes (ant-plants) are generally well protected from herbivore attacks by their symbiotic ants (plant-ants). However, larvae of Arhopala (Lepidoptera: Lycaenidae) species survive and develop on specific Macaranga ant-plant species without being attacked by the plant-ants of their host species. We hypothesized that Arhopala larvae chemically mimic or camouflage themselves with the ants on their host plant so that the larvae are accepted by the plant-ant species of their host. Chemical analyses of cuticular hydrocarbons showed that chemical congruency varied among Arhopala species; A. dajagaka matched well the host plant-ants, A. amphimuta did not match, and unexpectedly, A. zylda lacked hydrocarbons. Behaviorally, the larvae and dummies coated with cuticular chemicals of A. dajagaka were well attended by the plant-ants, especially by those of the host. A. amphimuta was often attacked by all plant-ants except for the host plant-ants toward the larvae, and those of A. zylda were ignored by all plant-ants. Our results suggested that conspicuous variations exist in the chemical strategies used by the myrmecophilous butterflies that allow them to avoid ant attack and be accepted by the plant-ant colonies.     

A castration parasite of an ant-plant mutualism

Exploring the factors governing the maintenance and breakdown of cooperation between mutualists is an intriguing and enduring problem for evolutionary ecology, and symbioses between ants and plants can provide useful experimental models for such studies. Hundreds of tropical plant species have evolved structures to house and feed ants, and these ant–plant symbioses have long been considered classic examples of mutualism. Here, we report that the primary ant symbiont, Allomerus cf. demerarae, of the most abundant ant-plant found in south-east Peru, Cordia nodosa Lam., castrates its host plant. Allomerus workers protect new leaves and their associated domatia from herbivory, but destroy flowers, reducing fruit production to zero in most host plants. Castrated plants occupied by Allomerus provide more domatia for their associated ants than plants occupied by three species of Azteca ants that do not castrate their hosts. Allomerus colonies in larger plants have higher fecundity. As a consequence, Allomerus appears to benefit from its castration behaviour, to the detriment of C. nodosa. The C. nodosa–ant system exhibits none of the retaliatory or filtering mechanisms shown to stabilize cheating in other cooperative systems, and appears to persist because some of the plants, albeit a small minority, are inhabited by the three species of truly mutualistic Azteca ants.     

Change in biomass of symbiotic ants throughout the ontogeny of a myrmecophyte, Macaranga beccariana (Euphorbiaceae)

Macaranga myrmecophytes (ant-plants) provide their partner symbiotic ants (plant-ants) with food bodies as their main food, and they are protected by the plant-ants from herbivores. The amount of resource allocated to food bodies determines the plant-ant colony size and consequently determines the intensity of ant defense (anti-herbivore defense by plant-ants). As constraints in resource allocation change as plants grow, the plant-ant colony size is hypothesized to change with the ontogenesis of Macaranga myrmecophyte. To determine the ontogenetic change in the relative size of the plant-ant colony, we measured the dry weights of the whole plant-ant colony and all of the aboveground parts of trees at various ontogenetic stages for a myrmecophytic species (Macaranga beccariana) in a Bornean lowland tropical rain forest. Ant biomass increased as plant biomass increased. However, the rate of increase gradually declined, and the ant biomass appeared to reach a ceiling once trees began to branch. The ant/plant biomass ratio consistently decreased as plant biomass increased, with the rate of decrease gradually accelerating. We infer that the ontogenetic reduction in ant/plant biomass ratio is caused by an ontogenetic change in resource allocation to food rewards for ants related to the physiological changes accompanying the beginning of branching.     

Pollination systems in pioneer trees of the genus Macaranga (Euphorbiaceae) in Malaysian rainforests

Many species of Macaranga (Euphorbiacae) are fast‐growing pioneer trees with an important role in early succession in south‐east Asian rainforests. Within the genus, diverse types of ant–plant associations exist and it has therefore been a model system for studying mutualistic interactions. Little information existed up to now, however, on its reproductive biology. Our comparative study in the genus Macaranga in Sundaland revealed specific flower characteristics and uncommon brood‐site pollination systems: enclosed inflorescence morphologies with narrow entrances strongly restrict the set of flower visitors in many species. Thysanoptera were the most abundant insects in 20 of the 26 investigated Macaranga species and, in three species, heteropteran adults and larvae were dominant. Both insect groups used the flower chambers as breeding sites and fed on nectar‐producing trichomes inside the bracteoles. Thrips as well as heteropterans are assumed to contribute to pollination. Different Macaranga sections were associated with different flower visitors, suggesting isolation by different pollinators. Thrips pollination and myrmecophyty often occurred in the same sections. The development of enclosed flowers might have facilitated tight ant–plant interactions and prevent ant–pollinator conflicts. However, the complex ecosystems in which the mutualistic systems evolved are rapidly changed with unknown consequences for these specific interactions. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 935–953.     

A mutualism's indirect costs: the most aggressive plant bodyguards also deter pollinators

Plant defenses against herbivores may be costly if they exclude mutualists. Here, I test the hypothesis that aggressive ant bodyguards of plants deter pollinators, and explore mechanisms by which Ferocactus wislizeni , an extrafloral nectary bearing cactus, limits conflicts between its pollinators and bodyguards. Flower visitation by ants and pollinating bees differed among plants tended by four different ant species. The ant species most rarely found in flowers showed the strongest aversion to F. wislizeni flower petals in laboratory assays, suggesting that those structures may include an ant-deterrent. Species-specific estimates of mean ant abundance within flowers and aggressiveness towards other arthropods were used to distinguish the relative threat of ant attack in flowers on plants tended by each ant species. Pollinator surveys in 2003 and 2004 demonstrated that bee visitation rates and the duration of flower occupation differed among plants with different ant associates, decreasing as the threat of ant attack increased. Flowers on plants tended by Solenopsis xyloni , the best ant bodyguard, were more dangerous than those on plants tended by three milder species, due to that ants' greater aggressiveness and abundance within flowers. These flowers were visited by pollinators least frequently and for less time per visit, and produced fruits with significantly lower total seed mass, fewer seeds, and lighter individual seeds, relative to fruits from similarly-sized plants tended by three other ant species. As a result, the best bodyguard may indirectly constrain plant reproduction in some settings. Conflicts between mutualistic guilds may be particularly common in generalized systems, where there is variation in partner quality and in the relative importance of the protection and pollination mutualisms.     

Floral visitation by the Argentine ant reduces pollinator visitation and seed set in the coast barrel cactus, Ferocactus viridescens

Mounting evidence indicates that trade-offs between plant defense and reproduction arise not only from resource allocation but also from interactions among mutualists. Indirect costs of plant defense by ants, for example, can outweigh benefits if ants deter pollinators. Plants can dissuade ants from occupying flowers, but such arrangements may break down when novel ant partners infiltrate mutualisms. Here, we examine how floral visitation by ants affects pollination services when the invasive Argentine ant (Linepithema humile) replaces a native ant species in a food-for-protection mutualism with the coast barrel cactus (Ferocactus viridescens), which, like certain other barrel cacti, produces extrafloral nectar. We compared the effects of floral visitation by the Argentine ant with those of the most prevalent native ant species (Crematogaster californica). Compared to C. californica, the Argentine ant was present in higher numbers in flowers. Cactus bees (Diadasia spp.), the key pollinators in this system, spent less time in flowers when cacti were occupied by the Argentine ant compared to when cacti were occupied by C. californica. Presumably as a consequence of decreased duration of floral visits by Diadasia, cacti occupied by L. humile set fewer seeds per fruit and produced fewer seeds overall compared to cacti occupied by C. californica. These data illustrate the importance of mutualist identity in cases where plants balance multiple mutualisms. Moreover, as habitats become increasingly infiltrated by introduced species, the loss of native mutualists and their replacement by non-native species may alter the shape of trade-offs between plant defense and reproduction.     

Pterandra pyroidea: a case of pollination shift within neotropical Malpighiaceae

Background and Aims

Most Neotropical species of Malpighiaceae produce floral fatty oils in calyx glands to attract pollinating oil-collecting bees, which depend on this resource for reproduction. This specialized type of pollination system tends to be lost in members of the family that occur outside the geographic distribution (e.g. Africa) of Neotropical oil-collecting bees. This study focused on the pollination ecology, chemical ecology and reproductive biology of an oil flower species, Pterandra pyroidea (Malpighiaceae) from the Brazilian Cerrado. Populations of this species consist of plants with oil-secreting (glandular) flowers, plants with non-oil-secreting flowers (eglandular) or a mix of both plant types. This study specifically aims to clarify the role of eglandular morphs in this species.

Methods

Data on pollinators were recorded by in situ observations. Breeding system experiments were conducted by isolating inflorescences and by enzymatic reactions. Floral resources, pollen and floral oils offered by this species were analysed by staining and a combination of various spectroscopic methods.

Key Results

Eglandular flowers of P. pyroidea do not act as mimics of their oil-producing conspecifics to attract pollinators. Instead, both oil-producing and oil-free flowers depend on pollen-collecting bees for reproduction, and their main pollinators are bumble-bees. Floral oils produced by glandular flowers are less complex than those described in closely related genera.

Conclusions

Eglandular flowers represent a shift in the pollination system in which oil is being lost and pollen is becoming the main reward of P. pyroidea flowers. Pollination shifts of this kind have hitherto not been demonstrated empirically within Neotropical Malpighiaceae and this species exhibits an unusual transition from a specialized towards a generalized pollination system in an area considered the hotspot of oil-collecting bee diversity in the Neotropics. Transitions of this type provide an opportunity to study ongoing evolutionary mechanisms that promote the persistence of species previously involved in specialized mutualistic relationships.     

Do aphids actively search for ant partners?

The aphid–ant mutualistic relationships are not necessarily obligate for neither partners but evidence is that such interactions provide them strong advantages in terms of global fitness. While it is largely assumed that ants actively search for their mutualistic partners namely using volatile cues; whether winged aphids (i.e., aphids’ most mobile form) are able to select ant‐frequented areas had not been investigated so far. Ant‐frequented sites would indeed offer several advantages for these aphids including a lower predation pressure through ant presence and enhanced chances of establishing mutuaslistic interactions with neighbor ant colonies. In the field, aphid colonies are often observed in higher densities around ant nests, which is probably linked to a better survival ensured by ants’ services. Nevertheless, this could also result from a preferential establishment of winged aphids in ant‐frequented areas. We tested this last hypothesis through different ethological assays and show that the facultative myrmecophilous black bean aphid, Aphis fabae L., does not orientate its search for a host plant preferentially toward ant‐frequented plants. However, our results suggest that ants reduce the number of winged aphids leaving the newly colonized plant. Thus, ants involved in facultative myrmecophilous interactions with aphids appear to contribute to structure aphid populations in the field by ensuring a better establishment and survival of newly established colonies rather than by inducing a deliberate plant selection by aphid partners based on the proximity of ant colonies.     

Oviposition by mutualistic seed-parasitic pollinators and its effects on annual fitness of single- and multi-flowered host plants

The balance of intimate relationships between plants and seed-eating pollinators can depend on pollinator behaviour in relation to floral characters, such as flower size and flower number. Here, we examined how pollinator oviposition in relation to these traits affected annual fitness (seed output) of single- and multi-flowered Trollius europaeus along altitudinal gradients in subarctic Sweden and the French Alps. Small flies (Chiastocheta spp.) pollinate T. europaeus and their larvae feed on developing seeds. Assuming that late flowers in multi-flowered plants attracted flies to the earliest flower on the same plant, we expected more eggs and higher seed predation in early multiple flowers than in single flowers. Relative seed predation would thereby increase with flower number. Both in Sweden and the Alps, more eggs were placed on large flowers. Early multiple flowers were slightly larger than single flowers, and about twice the size of secondary flowers. As a result, and possibly combined with the effects of secondary flowers, early multiple flowers attracted more ovipositing flies and experienced relatively higher seed predation. However, this did not generally result in higher seed predation of multi-flowered hosts. Multiple flowers had greater seed output than single flowers at all altitudes, also in the high alpine and subarctic sites, where single flowers were more abundant. We hypothesise that the distribution of multiple flowers generally is enforced by environmental factors, rather than by fly-host plant interactions, because only very rarely (in triple-flowered alpine plants) was seed predation disproportionate, and the relationship skewed to the disadvantage of the host. The outcome of the mutualistic interaction was often similar in alpine and subarctic populations, but the underlying factors were different. Subarctic flowers had high abortion and low predation rates, while alpine flowers experienced the reversed situation. The higher fly abundance in the Alps suggests a more intense mutualistic interaction than in Sweden. Despite varying ecological and environmental conditions at these sites, the mutualistic relationship was generally in balance. However, when it was unbalanced, this could be explained by fly behaviour in response to floral traits, and by differences in fly abundance. Received: 4 January 1999 / Accepted: 5 May 1999     

Size-dependent reproduction in Australian alpine Ranunculus

Abstract The effect of plant size on reproduction in four species of alpine Ranunculus (R. muelleri, R. dissectifolius, R. graniticola and R. niphophilus) was investigated in two sites over two seasons in the field on a total of 190 plants. The effects of plant size (number of leaves) and number of flowers on the number of anthers, ovules and seed per flower and per plant were determined. There was a positive relationship between several measures of reproduction and plant size in all four species, indicating that reproduction is size-dependent. All the results indicate that the main factor controlling the amount of seed produced by alpine Ranunculus is the size of the plant. Specifically, bigger plants produced more seed by producing more flowers, not by producing more ovules per flower, or higher seed set per flower. Correspondingly, bigger plants produced more anthers by producing more flowers, rather than by producing flowers with more anthers. The total number of seeds produced by a plant was directly proportional to plant size in the four species. Therefore, reproductive effort should not vary with plant size in the four species.     

Costs and benefits of ant attendance to the drepanosiphid aphid Tuberculatus quercicola

The defensive effects of ants against aphid predators have been well documented in the mutualistic relationship of aphids and their attending ants. However, it is not clear whether ant attendance has any direct effect on the aphids' growth and reproduction. Through field experiments, this study evaluates the benefits and, in particular, the costs of ant attendance to aphid colonies, focusing on the drepanosiphid aphid Tuberculatus quercicola which is associated with the Daimyo oak, Quercus dentata , and which is always attended by the red wood ant Formica yessensis . Ant attendance was clearly beneficial to the aphid; the exclusion of ants led to a significant increase in the extinction rate of aphid colonies. However, MANOVA and randomized block ANOVA indicated that in colonies continuously attended by ants, aphids had significantly smaller body size and produced a smaller number of embryos than in colonies isolated from ants when they were reared under homogeneous host conditions free from natural enemies. Thus, ant attendance had a negative influence on the growth and reproduction of the aphids, even though it contributed to the greater longevity of the aphid colonies. We hypothesize that ant-attended aphids are under intense selective pressures that act against aphid clones which fail to attract many ants, so that aphids have developed an adaptive mechanism to allocate a larger fraction of resources to the honeydew when they are requested to do so by the ants in order to ensure the ants' consistent visitation.     

Novel interactions of non-pollinating ants with pollinators and fruit consumers in a tropical forest

The tropical ants Ectatomma ruidum and E. tuberculatum (Formicidae) regularly patrol leaves, flowers, and fruits of the understory shrub, Psychotria limonensis (Rubiaceae), on Barro Colorado Island, Panama. Ant and pollinator exclusion experiments elucidated both positive and negative effects of ant attendance on plant reproductive success, including pollination, fruit set, fruit loss, and fruit removal. Ants did not pollinate flowers but did contribute to higher pollination success, probably by increasing the relocation frequency of winged pollinators and thus the rate of flower visitation. Ants also prevented fruit loss to herbivorous insects which were common during the early stages of fruit development. Thus, ant attendance strongly improved both pollination and fruit set whereby plants with ants set more fruit per flower and also lost fewer fruits during fruit maturation. In contrast, ants had a negative effect on the removal of ripe fruits by avian frugivores. Thus, ant attendance has a non-trivial influence on plant reproduction, this interaction being beneficial at some stages of the plant reproductive cycle and carrying costs at another stage. A tight ecological or co-evolved relationship between these Ectatomma spp. and P. limonensis is unlikely given that ant attendance of plants is detrimental to fruit removal. Received: 18 May 1998 / Accepted: 1 March 1999     

The reproductive biology of two understory plants in the Atlantic rain forest, Brazil

Outcrossing and sexual reproduction of most flowering plants depends on pollinators. Plant traits likely to be involved in pollinator attraction include flower color, shape, and size. Furthermore, plant or flower density and the temporal flowering pattern may have an effect on reproduction. In this study, we examine the pollination ecology, breeding system, female reproductive output, and germination of two tropical understory species, Stenostephanus lobeliiformis (Acanthaceae) and Besleria melancholica (Gesneriaceae), which differ in these traits. Pollinator observations revealed that the dense flowering S. lobeliiformis with pinkish flowers received a higher diversity of pollinators, but visitor frequency measured as visits per flower per hour was much less (0.1 h^?1) than that to B. melancholica, which has a smaller floral display of dull-colored flowers (1.5 h^?1). Pollination experiments revealed that S. lobeliiformis but not B. melancholica is pollen-limited. In addition, both species are partially self-incompatible and depend on pollinators for outcrossing. Natural fruit set of open-pollinated unmanipulated flowers (control treatment) in both species is 22–26 %. Germination studies indicated inbreeding depression in S. lobeliiformis. We conclude that the pollination ecology of these species is influenced by a broad set of traits and that very different combinations of these traits can be successful in terms of reproduction.     

Production of food bodies on the reproductive organs of myrmecophytic Macaranga species (Euphorbiaceae): effects on interactions with herbivores and pollinators

In protective ant–plant mutualisms, plants offer ants food (such as extrafloral nectar and/or food bodies) and ants protect plants from herbivores. However, ants often negatively affect plant reproduction by deterring pollinators. The aggressive protection that mutualistic ants provide to some myrmecophytes may enhance this negative effect in comparison to plant species that are facultatively protected by ants. Because little is known about the processes by which myrmecophytes are pollinated in the presence of ant guards, we examined ant interactions with herbivores and pollinators on plant reproductive organs. We examined eight myrmecophytic and three nonmyrmecophytic Macaranga species in Borneo. Most of the species studied are pollinated by thrips breeding in the inflorescences. Seven of eight myrmecophytic species produced food bodies on young inflorescences and/or immature fruits. Food body production was associated with increased ant abundance on inflorescences of the three species observed. The exclusion of ants from inflorescences of one species without food rewards resulted in increased herbivory damage. In contrast, ant exclusion had no effect on the number of pollinator thrips. The absence of thrips pollinator deterrence by ants may be due to the presence of protective bracteoles that limit ants, but not pollinators, from accessing flowers. This unique mechanism may account for simultaneous thrips pollination and ant defense of inflorescences.     

Dependency on floral resources determines the animals' responses to floral scents

Animal-pollinated angiosperms either depend on cross-pollination or may also reproduce after self-pollination—the former are thus obligately, the latter facultatively dependent on the service of animal-pollinators. Analogously, flower visitors either solely feed on floral resources or complement their diet with these, and are hence dependent or not on the flowers they visit. We assume that obligate flower visitors evolved abilities that enable them to effectively forage on flowers including mechanisms to bypass or tolerate floral defences such as morphological barriers and repellent/deterrent secondary metabolites. Facultative flower visitors, in contrast, are supposed to lack these adaptations and are often prevented to consume floral resources by defence mechanisms. In cases where obligate flower visitors are mutualists and facultative ones are antagonists, this dichotomy provides a solution for the plants'' dilemma to attract pollinators and simultaneously repel exploiters. In a meta-analysis, we recently supported this hypothesis: obligate flower visitors are attracted to floral scents, while facultative ones are repelled. Here, we add empirical evidence to these results: bumblebees and ants, obligate and facultative flower visitors, respectively, responded as predicted by the results of the meta-analysis to synthetic floral scent compounds.Key words: antagonists, exploitation, floral defences, mutualism, nectar, pollinationThe mutualism between flowers and their pollinators is often exploited by cheaters that consume floral rewards but do not contribute to or even reduce the reproductive success of plants.¹ The classification into mutualistic and antagonistic flower visitors represents a phytocentric point of view and only considers the interaction''s net effect for the plant. However, the outcome of each plant-flower visitor interaction may be highly conditional and variable over time² and thus constitutes a continuum between beneficial and detrimental, and it may not be unequivocally assigned to be either positive or negative. Furthermore, many flower visitor species that function as effective pollinators of some plant species represent severe antagonists to other plant species.³ Thus, except for highly specialised systems, it is difficult to predict whether an interaction is mutualistic, commensalistic or antagonistic. We proposed a different classification of flower visitors based on the animals'' interest in flower visits.⁴ Animals visit flowers primarily in search for food; pollination is just a secondary effect.⁵ For some taxa nectar and pollen are the sole nutrient supply, others only supplement their more generalistic diets with floral resources. These different dependencies on floral resources can often be unequivocally assigned to each animal species. Bees, for example, strongly depend on pollen and nectar and are thus obligate flower visitors. In contrast, ants are omnivores and thus facultative flower visitors that consume large amounts of floral nectar of some plant species but obtain most of the nutrients required by the colony from non-floral resources.⁶Optimal foraging theory predicts that animals evolve physiological and behavioral features that allow them to exploit their resources as effectively as possible.^7,8 Therefore, a classification considering the animals'' dependencies on floral resources (obligate versus facultative) may be better suited to explain adaptations to flower visits than their effect on plants'' reproduction (mutualistic versus antagonistic). One very important adaptation to the consumption of floral resources is the ability to tolerate or overcome floral defences that are employed by the flowers to reduce the visitation frequency of detrimental flower visitors.⁹ Floral scents are innate attractants or reinforce floral visits due to associative learning but do also serve as effective repellents against antagonists.¹⁰ In a meta-analysis, we recently demonstrated that the dependency on floral resources determines the responses to floral scents.⁴ In the bioassay presented here, using bumblebees (Bombus terrestris) and ants (Lasius niger), we empirically tested the predictions deduced from the metaanalysis. We expected that bumblebees—as obligate flower visitors—are attracted to floral scent compounds, while ants—as facultative flower visitors—are repelled.     

Anthropogenic edges,isolation and the flowering time and fruit set of Anadenanthera peregrina,a cerrado savanna tree

Fragmentation exposes plants to extreme environmental conditions with implications for species phenology and reproduction. We investigated whether isolation and edge effects influence size, flowering time, fruit set, and seedling establishment of Anadenanthera peregrina var. falcata. We compared trees in the interior (n?=?85), and on the edge (n?=?74) of a cerrado savanna fragment as well as in a pasture (n?=?26) with respect to size, flowering phenology, flower and fruit production, fruit and seed set, predispersal seed predation, and seedling establishment. Trees in the pasture were larger and produced a higher number of flowers and fruits than trees on the edge and interior, yet seed set did not differ across environments. The plant size structure explained the flower and fruit production, and the self-compatibility breeding system caused a similar seed set regardless of the environment. First flowering was later and fruit set higher in the interior. We argue that time of first flower influenced the fruit set of Anadenathera. Edge and isolated trees started to flower earlier as a response to microclimatic conditions—mainly temperature—reducing the fruit set. Predispersal seed predation was lower among pasture trees. Conversely, we found seedlings only on the edge and in the interior of cerrado, suggesting that the pasture was of poor quality habitat for Anadenanthera recruitment. Isolation affected the plant size structure and reproduction of Anadenanthera trees. Studies comparing plant phenology under contrasting environmental conditions may offer clues on how global change may affect plant reproduction in the tropics.     

Pollination by ants: consequences of the quantitative effects on a mutualistic system

Summary We have analysed the importance of worker ants (Proformica longiseta, Formicidae) as pollinators of a mass-flowering woody plant (Hormathophylla spinosa, Cruciferae) in the high-mountain area of the Sierra Nevada (southern Spain). We have quantified the abundance and foraging behavior of P. longiseta in comparison with winged flower visitors. We have also examined, by means of selective exclusion experiments, the role of ants as true pollinators, comparing them with the winged flower visitors. A total of 39 species belonging to 18 families visited the flowers of H. spinosa. All the visitors were winged insects, except P. longiseta, a species which alone made up more than 80% of the total number of insects found on the flowers. All pollinators of H. spinosa had similar foraging patterns, with 98% of total movements made between flowers within the same plant. Ants always made contact with the plant reproductive organs when foraging for nectar, and transferred large numbers of pollen grains. However, pollen exposed to ants for brief periods exhibited reduced percentage of germination. P. longiseta is both the most abundant and spatio-temporally predictable flower visitor of H. spinosa. These characteristics, weighted by their flower visitation rate, make worker ants the pollinator that maintains the strongest mutualistic interaction with H. spinosa. The exclusion experiments show that workers behave as true pollinators, since they contribute to increase the number of viable seeds produced by H. spinosa. The key factor of this interaction is mainly the great density of workers throughout the flowering period. In short, the H. spinosa-P. longiseta mutualistic interaction mainly depends on its high probability of occurrence.