The ontogeny of plant indirect defenses

Plants frequently attract natural enemies of their herbivores, resulting in a reduction in tissue damage and often in enhanced plant fitness. While such indirect defenses can dramatically change as plants develop, only recently have ecologists begun to explore such changes and evaluate their role in mediating plant–herbivore–natural enemy interactions. Here we review the literature documenting ontogenetic patterns in plant rewards (i.e. extrafloral nectaries (EFNs), food bodies (FBs) and domatia) and volatile organic compounds (VOCs), and identify links between ontogenetic patterns in such traits and the attraction of natural enemies (ants). In the case of reward traits we concentrate in ant–plant studies, which are the most numerous. We report that all indirect defensive traits commonly vary with plant age but ontogenetic trajectories differ among them. Myrmecophytic species, which provide both food and shelter to their defenders, do not produce rewarding traits until a minimum size is reached. Then, a pronounced increase in the abundance of food rewards and domatia often occurs as plants develop, which explains the temporal succession or colony size increase of mutualistic ant species and, in some cases, leads to a reduction in herbivore damage and enhanced fitness as plants age. In contrast, ontogenetic patterns were less consistent in plant species that rely on VOC emissions to attract natural enemies or those that provide only food rewards (EFNs) but not nesting sites to their associated ants, showing an overall decline or lack of trend with plant development, respectively. Future research should focus on uncovering: (i) the costs and mechanisms underlying ontogenetic variation in indirect defenses, (ii) the relative importance of environmental and genetic components shaping these ontogenetic trajectories, and (iii) the consequences of these ontogenetic trajectories on plant fitness. Advances in this area will shed light on the context dependency of bottom-up and top-down controls of herbivore populations and on how natural selection actually shapes the ontogenetic trajectories of these traits.     

Natural selection on extrafloral nectar production in Chamaecrista fasciculata: the costs and benefits of a mutualism trait

Cost-benefit models of the evolution of mutualism predict that the current state of mutualism results from trade-offs between fitness costs of mutualist traits and the fitness benefits of association. We test the assumptions of such models by measuring patterns of natural selection on a mutualist trait, extrafloral nectar production in Chamaecrista fasciculata. Selection was measured on plants from which ants had been excluded (removing the mutualist benefit of the trait), from which all insects had been excluded (removing costs of herbivory in addition to mutualist benefits), and unmanipulated plants (where both costs and benefits were present). Selection analysis based on half-sibling-mean regressions of fitness on the trait revealed no evidence of costs of extrafloral nectar production in the absence of all insects or in the absence of ants. However, examination of the selective surfaces for these treatments suggest that costs of nectar production may exist and are exacerbated by the presence of herbivory. In the presence of ants, natural selection favors high extrafloral nectar production, consistent with a fitness benefit to this mutualist trait in the presence of the mutualist partner. In this study, the interaction of costs and benefits did not produce an evolutionary optimum for the trait within the range of variation observed, suggesting that application of a cost-benefit framework to this trait will benefit from considering the influence of temporal and spatial variation on the quality of costs and benefits.     

The effect of disturbance on an ant–plant mutualism

Protective ant–plant mutualisms—where plants provide food or shelter to ants and ants protect the plants from herbivores—are a common feature in many ecological communities, but few studies have examined the effect of disturbance on these interactions. Disturbance may affect the relationship between plants and their associated ant mutualists by increasing the plants’ susceptibility to herbivores, changing the amount of reward provided for the ants, and altering the abundance of ants and other predators. Pruning was used to simulate the damage to buttonwood mangrove (Conocarpus erectus) caused by hurricanes. Pruned plants grew faster than unpruned plants, produced lower levels of physical anti-herbivore defenses (trichomes, toughness), and higher levels of chemical defenses (tannins) and extrafloral nectaries. Thus, simulated hurricane damage increased plant growth and the amount of reward provided to ant mutualists, but did not have consistent effects on other anti-herbivore defenses. Both herbivores and ants increased in abundance on pruned plants, indicating that the effects of simulated hurricane damage on plant traits were propagated to higher trophic levels. Ant-exclusion led to higher leaf damage on both pruned and upruned plants. The effect of ant-exclusion did not differ between pruned and unpruned plants, despite the fact that pruned plants had higher ant and herbivore densities, produced more extrafloral nectaries, and had fewer physical defenses. Another common predator, clubionid spiders, increased in abundance on pruned plants from which ants had been excluded. I suggest that compensatory predation by these spiders diminished the effect of ant-exclusion on pruned plants.     

Biotic resistance and invasional meltdown: consequences of acquired interspecific interactions for an invasive orchid,Spathoglottis plicata in Puerto Rico

Invasiveness of non-native species often depends on acquired interactions with either native or naturalized species. A natural colonizer, the autogamous, invasive orchid Spathoglottis plicata has acquired at least three interspecific interactions in Puerto Rico: a mycorrhizal fungus essential for seed germination and early development; a native, orchid-specialist weevil, Stethobaris polita, which eats perianth parts and oviposits in developing fruits; and ants, primarily invasive Solenopsis invicta, that forage at extrafloral nectaries. We tested in field experiments and from observational data whether weevils affect reproductive success in the orchid; and whether this interaction is density-dependent. We also examined the effectiveness of extrafloral nectaries in attracting ants that ward off weevils. Only at small spatial scales were weevil abundance and flower damage correlated with flower densities. Plants protected from weevils had less floral damage and higher fruit set than those accessible to weevils. The more abundant ants were on inflorescences, the less accessible fruits were to weevils, resulting in reduced fruit loss from larval infections. Ants did not exclude weevils, but they affected weevil activity. Native herbivores generally provide some biotic resistance to plant invasions yet Spathoglottis plicata remains an aggressive colonizer despite the acquisition of a herbivore/seed predator partly because invasive ants attracted to extrafloral nectaries inhibited weevil behavior. Thus, the invasion of one species facilitates the success of another as in invasional meltdowns. For invasive plant species of disturbed habitats, having ant-tended extrafloral nectaries and producing copious quantities of seed, biotic resistance to plant invasions can be minimal.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

How do plants balance multiple mutualists? Correlations among traits for attracting protective bodyguards and pollinators in cotton (Gossypium)

Many species, both plants and animals, are simultaneously engaged in interactions with multiple mutualists. However, the extent to which separate traits that attract different mutualist guilds display negative or positive relationships remains largely unstudied. We asked whether correlations exist among extrafloral nectary traits to attract arthropod bodyguards and floral traits to attract pollinator mutualists. For 37 species in the cotton genus (Gossypium), we evaluated correlations among six extrafloral nectary traits and four floral traits in a common greenhouse environment, with and without correction for phylogenetic non-independence. Across Gossypium species, greater investment in extrafloral nectary traits was positively correlated with greater investment in floral traits. Positive correlations remained after accounting for the evolutionary history of the clade. Our results demonstrate that traits to maintain multiple mutualist guilds can be positively correlated across related species and build a more general understanding of the constraints on trait evolution in plants.     

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.     

Geographic variation in a facultative mutualism: consequences for local arthropod composition and diversity

Geographic variation in the outcome of interspecific interactions may influence not only the evolutionary trajectories of species but also the structure of local communities. We investigated this community consequence of geographic variation for a facultative mutualism between ants and wild cotton (Gossypium thurberi). Ants consume wild cotton extrafloral nectar and can protect plants from herbivores. We chose three sites that differed in interaction outcome, including a mutualism (ants provided the greatest benefits to plant fitness and responded to manipulations of extrafloral nectar), a potential commensalism (ants increased plant fitness but were unresponsive to extrafloral nectar), and a neutral interaction (ants neither affected plant fitness nor responded to extrafloral nectar). At all sites, we manipulated ants and extrafloral nectar in a factorial design and monitored the abundance, diversity, and composition of other arthropods occurring on wild cotton plants. We predicted that the effects of ants and extrafloral nectar on arthropods would be largest in the location with the mutualism and weakest where the interaction was neutral. A non-metric multidimensional scaling analysis revealed that the presence of ants altered arthropod composition, but only at the two sites in which ants increased plant fitness. At the site with the mutualism, ants also suppressed detritivore/scavenger abundance and increased aphids. The presence of extrafloral nectar increased arthropod abundance where mutual benefits were the strongest, whereas both arthropod abundance and morphospecies richness declined with extrafloral nectar availability at the site with the weakest ant–plant interaction. Some responses were geographically invariable: total arthropod richness and evenness declined by approximately 20% on plants with ants, and extrafloral nectar reduced carnivore abundance when ants were excluded from plants. These results demonstrate that a facultative ant–plant mutualism can alter the composition of arthropod assemblages on plants and that these community-level consequences vary across the landscape.     

Phylogenetic and Experimental Tests of Interactions among Mutualistic Plant Defense Traits in Viburnum (Adoxaceae)

Abstract Plant traits that mediate mutualistic interactions are widespread, yet few studies have linked their macroevolutionary patterns with the ecological interactions they mediate. Here we merged phylogenetic and experimental approaches to investigate the evolution of two common mutualistic plant traits, extrafloral nectaries (EFNs) and leaf domatia. By using the flowering plant clade Viburnum, we tested whether macroevolutionary patterns support adaptive hypotheses and conducted field surveys and manipulative experiments to examine whether ecological interactions are concordant with evolutionary predictions. Phylogenetic reconstructions suggested that EFN-bearing species are monophyletic, whereas the evolution of domatia correlated with leaf production strategy (deciduous or evergreen) and climate. Domatia were also more common in the EFN clade, suggesting that the two traits may jointly mediate ecological interactions. This result was further investigated in a common-garden survey, where plants with domatia and EFNs on the leaf blade had more mutualistic mites than plants with other trait combinations, and in manipulative field experiments, where the traits additively increased mutualist abundance. Taken together, our results suggest that mutualistic traits in Viburnum are not ecologically independent, as they work in concert to attract and retain mutualists, and their long-term evolution may be influenced by complex interactions among multiple traits, mutualists, and geography.     

Evolution of extrafloral nectaries: adaptive process and selective regime changes from forest to savanna

Much effort has been devoted to understanding the function of extrafloral nectaries (EFNs) for ant–plant–herbivore interactions. However, the pattern of evolution of such structures throughout the history of plant lineages remains unexplored. In this study, we used empirical knowledge on plant defences mediated by ants as a theoretical framework to test specific hypotheses about the adaptive role of EFNs during plant evolution. Emphasis was given to different processes (neutral or adaptive) and factors (habitat change and trade‐offs with new trichomes) that may have affected the evolution of ant–plant associations. We measured seven EFN quantitative traits in all 105 species included in a well‐supported phylogeny of the tribe Bignonieae (Bignoniaceae) and collected field data on ant–EFN interactions in 32 species. We identified a positive association between ant visitation (a surrogate of ant guarding) and the abundance of EFNs in vegetative plant parts and rejected the hypothesis of phylogenetic conservatism of EFNs, with most traits presenting K‐values < 1. Modelling the evolution of EFN traits using maximum likelihood approaches further suggested adaptive evolution, with static‐optimum models showing a better fit than purely drift models. In addition, the abundance of EFNs was associated with habitat shifts (with a decrease in the abundance of EFNs from forest to savannas), and a potential trade‐off was detected between the abundance of EFNs and estipitate glandular trichomes (i.e. trichomes with sticky secretion). These evolutionary associations suggest divergent selection between species as well as explains K‐values < 1. Experimental studies with multiple lineages of forest and savanna taxa may improve our understanding of the role of nectaries in plants. Overall, our results suggest that the evolution of EFNs was likely associated with the adaptive process which probably played an important role in the diversification of this plant group.     

Ants impact sawfly oviposition on bracken fern in southern California

Although it has been speculated that ant visits to extrafloral nectaries of bracken fern may convey a fitness benefit for the plant, this has never been demonstrated with native herbivores and natural insect densities. We tested the hypothesis that ants attracted to extrafloral nectaries of bracken fern provide a mutualistic benefit by protecting fronds from herbivore damage in a field manipulation experiment in southern California. We examined densities of sawfly eggs and larvae on bracken fronds with and without ant exclusion. Because bracken fern in this region is also impacted by nitrogenous air pollution, we included an N addition treatment. We found that sawfly egg abundance was significantly higher for fern plants when ants were excluded, regardless of N treatment. Ants tended to have higher abundance on fertilized plants, but there was no interaction between N additions and ant exclusion. Bracken fern may derive a fitness benefit from attracting ants during the early phases of plant growth, through decreased herbivore oviposition, rather than through the deterrence of feeding larvae.     

Extrafloral nectaries in the genus Macaranga (Euphorbiaceae) in Malaysia: comparative studies of their possible significance as predispositions for myrmecophytism

Some species of the paleotropical tree genus Macaranga (Euphorbiaceae) live in close association with ants. The genus comprises the full range of species from those not regularly inhabited by ants to obligate myrmecophytes. In Malaysia (Peninsular and Borneo) 23 of the 52 species are known to be ant-associated (44%). The simplest structural adaptation of plants to attract ants are extrafloral nectaries. We studied the distribution of extrafloral nectaries in the genus Macaranga to assess the significance of this character as a possible predisposition for the evolution of obligate myrmecophytism. All species have marginal glands on the leaves. However, only the glands of non- myrmecophytic species function as nectaries, whereas liquids secreted by these glands in myrmecophytic species did not contain sugar. Some non-myrmecophytic Macaranga and transitional Macaranga species in addition have extrafloral nectaries on the leaf blade near the petiole insertion. All obligatorily myrmecophytic Macaranga species, however, lack additional glands on the lamina. The non-myrmecophytic species are visited by a variety of different ant species, whereas myrmecophytic Macaranga are associated only with one specific ant-partner. Since these ants keep scale insects in the hollow stems, reduction of nectary production in ant-inhabited Macaranga seems to be biologically significant. We interpret this as a means of (a) saving the assimilates and (b) stabilization of maintenance of the association's specificity. Competition with other ant species for food rewards is avoided and thereby danger of weakening the protective function of the obligate ant- partner for the plant is reduced. A comparison with other euphorb species living in the same habitats as Macaranga showed that in genera in which extrafloral nectaries are widespread, no myrmecophytes have evolved. Possession of extrafloral nectaries does not appear to be essential for the development of symbiotic ant-plant interactions. Other predispositions such as nesting space might have played a more important role.     

Geographic Mosaic of Plant Evolution: Extrafloral Nectary Variation Mediated by Ant and Herbivore Assemblages

Herbivory is an ecological process that is known to generate different patterns of selection on defensive plant traits across populations. Studies on this topic could greatly benefit from the general framework of the Geographic Mosaic Theory of Coevolution (GMT). Here, we hypothesize that herbivory represents a strong pressure for extrafloral nectary (EFN) bearing plants, with differences in herbivore and ant visitor assemblages leading to different evolutionary pressures among localities and ultimately to differences in EFN abundance and function. In this study, we investigate this hypothesis by analyzing 10 populations of Anemopaegma album (30 individuals per population) distributed through ca. 600 km of Neotropical savanna and covering most of the geographic range of this plant species. A common garden experiment revealed a phenotypic differentiation in EFN abundance, in which field and experimental plants showed a similar pattern of EFN variation among populations. We also did not find significant correlations between EFN traits and ant abundance, herbivory and plant performance across localities. Instead, a more complex pattern of ant–EFN variation, a geographic mosaic, emerged throughout the geographical range of A. album. We modeled the functional relationship between EFNs and ant traits across ant species and extended this phenotypic interface to characterize local situations of phenotypic matching and mismatching at the population level. Two distinct types of phenotypic matching emerged throughout populations: (1) a population with smaller ants (Crematogaster crinosa) matched with low abundance of EFNs; and (2) seven populations with bigger ants (Camponotus species) matched with higher EFN abundances. Three matched populations showed the highest plant performance and narrower variance of EFN abundance, representing potential plant evolutionary hotspots. Cases of mismatched and matched populations with the lowest performance were associated with abundant and highly detrimental herbivores. Our findings provide insights on the ecology and evolution of plant–ant guarding systems, and suggest new directions to research on facultative mutualistic interactions at wide geographic scales.     

Variation in the composition and activity of ants on defense of host plant <Emphasis Type="Italic">Turnera subulata</Emphasis> (Turneraceae): strong response to simulated herbivore attacks and to herbivore’s baits

Plants with extrafloral nectaries attract a variety of ant species, in associations commonly considered mutualistic. However, the results of such interactions can be context dependent. Turnera subulata is a shrub widely distributed among disturbed areas which has extrafloral nectaries at the base of leaves. Here, we evaluated whether the ants associated with T. subulata (i) vary in space and/or time; (ii) respond to simulated herbivory, and (iii) reduce herbivory rates. For this, we quantified the abundance and species richness of ants associated with T. subulata throughout the day in six different sites and the defensive capability of these ants under simulated herbivory in the leaves and stems of T. subulata plants (N = 60). We also checked the proportion of the lost leaf area and quantified leaf damage by chewing herbivores in the host plant. We found that a total of 21 ant species associated with the host plant. Species composition showed significant variation across the sampled sites and throughout the day. Visitation rates and predation by ants were higher in plant stems than in leaves. In general, herbivory rates were not correlated with ant association or activity, with the exception of the proportion of leaf area consumed; there was a significant lower herbivory rate on plants in which ants defended the leaves. Our results suggest that the benefits of association may depend on the ecological context. This context dependence may mask the correlation between the defense of ants and herbivory rates.     

Insights into a novel three-partner interaction between ants,coreids (Hemiptera: Coreidae) and extrafloral nectaries: implications for the study of protective mutualisms

Extrafloral nectar of plants and honeydew of hemipterans are the common mediators of facultative interactions that involve ants as a mobile strategy of defence. The outcome of these interactions can vary from mutualistic to commensalistic or even antagonistic, depending on the ecological context and the interacting species. Here, we explore a novel, three-partner interaction involving ants, the coreid Dersagrena subfoveolata (Hemiptera) and the extrafloral nectaries (EFNs) bearing plant Senna aphylla (Fabaceae) in semi-arid Northwest Argentina. We surveyed natural areas and conducted ant exclusion experiments, to understand how each pairwise interaction influences the overall outcome among the three interacting parts. The outcome of the interactions was assessed for experimental plants as the reproductive output and herbivore abundances and for coreids as predator abundances. We found that the coreids occurred exclusively on S. aphylla plants and that at least nine ant species interacted with the EFNs as well as with the coreids. Coreid occurrence and abundance depended on ant densities, which in turn, was determined by the presence of actively secreting EFNs. Coreid and ant presence did not influence plant reproductive success, and ants provided to coreids some biotic defence, mainly against vespid wasp predators, but had no effect on non-coreid herbivores. We conclude that the interaction outcome is commensalistic between ants and plants (assuming that EF nectar is not costly for the plant), antagonistic between coreids and plants, and mutualistic between coreids and ants. The sum of all outcomes is net positive effect for ants and coreids, and net slightly negative to neutral for plants.     

No Defensive Role of Ants throughout a Broad Latitudinal and Elevational Range of a Cactus

Defensive mutualisms mediated by extrafloral nectaries are particularly variable; their net results may change with seasons, communities and environmental contexts. Particularly, an environmental factor that can promote changes in outcomes of ant‐plant interactions is elevation in mountainous regions. We tested whether (1) the interaction between the cactus Opuntia sulphurea and ant visitors of extrafloral nectaries is a defensive mutualism; and (2) ant‐plant interaction outcomes vary with elevation as a result of changes in herbivory rate and ant activity. To evaluate if the outcome of interactions was consistent at two extremes of the range distribution of O. sulphurea, we performed an ant‐exclusion experiment with plants at two growth conditions (natural or potted) in two sites with contrasting elevation (1235–1787 m asl) in a temperate region (Villavicencio Nature Reserve, Mendoza, Argentina), and in a tropical region (Huajchilla, La Paz, Bolivia). Although herbivory rate and ant visitation frequency increased with elevation, herbivore damage, plant reproductive success, and cladode growth rate were similar between plants excluded and non‐excluded from ants among sites, geographic regions and growth conditions. These results do not support the hypotheses that the interaction between O. sulphurea and ants is a defensive mutualism, and that elevation affects the net outcome of this ant‐plant interaction.     

Disturbance Winners or Losers? Plants Bearing Extrafloral Nectaries in Brazilian Caatinga

Plants bearing extrafloral nectaries (EFNs) often have traits typical of pioneer species, and may be expected to proliferate in disturbed habitats. However, a negative effect of disturbance on visitation by attendant ants could prevent EFN‐bearing plants from acting as disturbance winners. Here, we address the effects of chronic anthropogenic disturbance on the abundance of EFN‐bearing plants and their interactions with attendant ants in Caatinga vegetation of northeastern Brazil. We recorded the abundance of EFN‐bearing plants, proportion of plants visited by ants and composition of ant attendant species at 24 sites varying in levels of disturbance. EFN‐bearing plants as a whole did not behave as a disturbance winner group. The responses of the 13 species to increasing disturbance were highly variable, with three species declining in abundance (loser species). The richness of ant species attending EFNs did not vary with disturbance, but species composition did. The overall proportion of EFN‐bearing plants attended by ants per 5‐min period was not affected by disturbance. However, for the three loser species, attendance decreased from about 50 percent with low and moderate disturbance to half that with very high disturbance. We hypothesize that disturbed sites are more stressful for loser species compared with other EFN‐bearing plants, with physiological stress resulting in lower production of EFN secretions and reduced attraction of ants. This would make such species double losers, with physiological stress at disturbed sites not only directly influencing their performance but also indirectly affecting it through the disruption of a key mutualism.     

Nectar quality affects ant aggressiveness and biotic defense provided to plants

Ant–plant mutualisms are useful models for investigating how plant traits mediate interspecific interactions. As plant‐derived resources are essential components of ant diets, plants that offer more nutritious food to ants should be better defended in return, as a result of more aggressive behavior toward natural enemies. We tested this hypothesis in a field experiment by adding artificial nectaries to individuals of the species Vochysia elliptica (Vochysiaceae). Ants were offered one of four liquid foods of different nutritional quality: amino acids, sugar, sugar + amino acids, and water (control). We used live termites (Nasutitermes coxipoensis) as herbivore competitors and observed ant behavior toward them. In 88 hr of observations, we recorded 1,009 interactions with artificial nectaries involving 1,923 individual ants of 26 species. We recorded 381 encounters between ants and termites, of which 38% led to attack. Sixty‐one percent of these attacks led to termite exclusion from the plants. Recruitment and patrolling were highest when ants fed upon nectaries providing sugar + amino acids, the most nutritious food. This increase in recruitment and patrolling led to higher encounter rates between ants and termites, more frequent attacks, and faster and more complete termite removal. Our results are consistent with the hypothesis that plant biotic defense is mediated by resource quality. We highlight the importance of qualitative differences in nectar composition for the outcome of ant–plant interactions. Abstract in Portuguese is available with online material.     

Resistance Mechanisms Against Arthropod Herbivores in Cotton and Their Interactions with Natural Enemies

Cotton plants (genus Gossypium) are grown on more than 30 million hectares worldwide and are a major source of fiber. The plants possess a wide range of direct and indirect resistance mechanisms against herbivorous arthropods. Direct resistance mechanisms include morphological traits such as trichomes and a range of secondary metabolites. The best known insecticidal compounds are the terpenoid gossypol and its precursors and related compounds. Indirect resistance mechanisms include herbivore-induced volatiles and extrafloral nectaries that allow plants to attract and sustain natural enemy populations. We discuss these resistance traits of cotton, their induction by herbivores, and their impact on herbivores and natural enemies. In addition, we discuss the use of genetically engineered cotton plants to control pest Lepidoptera and the influence of environmental factors on the resistance traits.     

Contrasting exotic Solenopsis invicta and native Forelius pruinosus ants as mutualists with Catalpa bignonioides, a native plant

Abstract. 1. The suitability of the red imported fire ant Solenopsis invicta Buren and a native ant Forelius pruinosus (Roger) as participants in a food-for-protection mutualism with a native nectaried tree Catalpa bignonioides Walter was compared.
2. The mean mortality of folivore larvae of Ceratomia catalpae Boisduval was similar for S. invicta and F. pruinosus although S. invicta attacked fewer caterpillar aggregations and was a devastating pupal predator. Solenopsis invicta also differed from the native ant in that it attacked the parasitoid Cotesia congregata Say, another plant mutualist, and visited extrafloral nectaries less frequently.
3. Habitats invaded by S. invicta are characterised by a scarcity of both herbivores and of beneficial insects that visit extrafloral nectaries. The plants do not require protection, and extrafloral nectaries are visited rarely. Although plants are defended incidentally by S. invicta , the insect-plant mutualism therein is greatly simplified or defunct.