Targeted predation of extrafloral nectaries by insects despite localized chemical defences

Extrafloral (EF) nectaries recruit carnivorous arthropods that protect plants from herbivory, but they can also be exploited by nectar thieves. We studied the opportunistic, targeted predation (and destruction) of EF nectaries by insects, and the localized chemical defences that plants presumably use to minimize this effect. In field and laboratory experiments, we identified insects that were possibly responsible for EF nectary predation in Vicia faba (fava bean) and determined the extent and accuracy of the feeding damage done to the EF nectaries by these insects. We also performed biochemical analyses of plant tissue samples in order to detect microscale distribution patterns of chemical defences in the area of the EF nectary. We observed selective, targeted feeding on EF nectaries by several insect species, including some that are otherwise not primarily herbivorous. Biochemical analyses revealed high concentrations of l-3,4-dihydroxyphenylalanine, a non-protein amino acid that is toxic to insects, near and within the EF nectaries. These results suggest that plants allocate defences to the protection of EF nectaries from predation, consistent with expectations of optimal defence theory, and that this may not be entirely effective, as insects limit their exposure to these defences by consuming only the secreting tissue of the nectary.     

Ant associations in the Neotropical shrub Turnera subulata (Turneraceae): Costs or benefits to the host plant?

The results of ecological interactions depend on the costs and benefits involved in different ecological contexts. Turnera subulata is a shrubby plant with extrafloral nectaries that are associated with ants. Here, we test the hypotheses that the association between T. subulata and ants results in: (i) positive effects on host plant growth and reproduction; (ii) plant herbivory reduction and (iii) inhibition of the host plant visitation by beneficial organisms. Thirty experimental plots were established in northeastern Brazil, either in association with ants or without ants (N = 15 plots/treatment), with four plants each (total 120 plants). Vegetative growth (plant height and number of leaves), reproductive investment (flowers and fruits), herbivory rates and numbers of beneficial visitors were quantified during all phenological stages of the host plant. Data were analysed using generalized linear mixed models. At the host plant maturation stage, we found a trade‐off between growth and reproduction. Plants with ants had lower mean height; however, they invested more in reproduction (a higher number of flowers and fruits) compared to plants without ants. During the flowering stage, the abundance of sucking herbivores was higher in plots without ants but chewing herbivore abundance increased in the maturation stage in plots with ants. The cumulative proportion of leaves with herbivore damage did not differ between treatments, and the presence of ants reduced the number of beneficial visitors (e.g. pollinators and natural enemies) to the host plants. Our results show that association with ants results in some costs for the host plant, however, these costs appear to be offset by the defensive role of ants, which favours plant reproductive investment. In general, our results help to elucidate mechanisms involved in trophic interactions within the complex network of interactions involving ants and plants.     

How common is the ability of extrafloral nectaries to produce nectar droplets,to secrete nectar during the night and to store starch?

Evidence in favour of the ability of extrafloral nectaries (EFNs) to form nectar drop(let)s, secrete extrafloral nectar (EFNec) also during the night and store starch was compiled in order to refute controversial assertions. Not only were more than 150 reports of direct observations of EFNec drop(let)s found, but also 90 studies which suggest that EFNec secretion is copious enough to form drop(let)s automatically by forces of physics (surface tension strength), provided nectar accumulation is not interrupted by predatory animals. Twenty direct observations of nocturnal production of EFNec sufficiently proved that it is not always produced during the day. Additionally, numerous observations of the nocturnal activities of nectar consumers on EFNs indirectly indicated very common nocturnal secretion of EFNec. Although there is an early report of a starch‐containing EFN from 1881 (Trelease), few similar observations in other EFNs followed. Nevertheless, four studies have described the disappearance of stored starch during secretion and senescence of the EFNs. Referring back to an apparent relationship between the degradation of starch stored in a floral nectary and programmed cell death, at least in EFNs with transient storage of starch, a similar relationship cannot be excluded.     

The diversity,ecology and evolution of extrafloral nectaries: current perspectives and future challenges

Background

Plants in over one hundred families in habitats worldwide bear extrafloral nectaries (EFNs). EFNs display a remarkable diversity of evolutionary origins, as well as diverse morphology and location on the plant. They secrete extrafloral nectar, a carbohydrate-rich food that attracts ants and other arthropods, many of which protect the plant in return. By fostering ecologically important protective mutualisms, EFNs play a significant role in structuring both plant and animal communities. And yet researchers are only now beginning to appreciate their importance and the range of ecological, evolutionary and morphological diversity that EFNs exhibit.

Scope

This Highlight features a series of papers that illustrate some of the newest directions in the study of EFNs. Here, we introduce this set of papers by providing an overview of current understanding and new insights on EFN diversity, ecology and evolution. We highlight major gaps in our current knowledge, and outline future research directions.

Conclusions

Our understanding of the roles EFNs play in plant biology is being revolutionized with the use of new tools from developmental biology and genomics, new modes of analysis allowing hypothesis-testing in large-scale phylogenetic frameworks, and new levels of inquiry extending to community-scale interaction networks. But many central questions remain unanswered; indeed, many have not yet been asked. Thus, the EFN puzzle remains an intriguing challenge for the future.     

Geographical and Seasonal Variation in the Richness of Ant-Plant Interactions in México1

The richness and seasonal variation of ant-plant interactions were compared in four habitats in México: lowland tropical dry forest (La Mancha, Veracruz), coastal sand dune matorral (San Benito, Yucatán), semiarid highland vegetation (Zapotitlán, Puebla), and lower montane humid forest (Xalapa, Veracruz). The effects of temperature and precipitation on the seasonal distribution of the number of ant-plant interactions differed among habitats. The general linear models fitted to the ant-plant interaction curves explained 78.8 percent of the variation for La Mancha, 80.1 percent for Zapotitlán, 18 percent for San Benito, and 29.5 percent for Xalapa. Even though rainfall is low in Zapotitlán and San Benito, minimum temperature was the most important factor accounting for the seasonal distribution and low number of interactions. At La Mancha, with milder minimum temperatures and higher water availability, temperature alone did not account for the seasonal distribution and number of interactions, whereas the effect of the precipitation × temperature interaction was highly significant. Xalapa exhibits the lowest temperatures and the highest precipitation, but the role of these factors was only marginal. We suggest that the vegetation at Xalapa, a mixture of tropical and temperate floristic elements, constrains ant-plant interactions due to a limited presence of nectaries. Also, ants are less abundant in cool and relatively aseasonal habitats. The other habitats have tropical floristic elements that are abundant and frequently have nectar-producing structures. We report considerable variation among habitats in the number and seasonal distribution of ant-plant interactions, and suggest that it is due to the effect of variation in environmental parameters, the richness of plants with nectaries in the vegetation, and habitat heterogeneity.     

Distribution and abundance of plants with extrafloral nectaries in the woody flora of a lowland primary forest in Malaysia

The first data on the taxonomic distribution and abundance of woody plants with extrafloral nectaries (EFN) from SE Asia are reported. The species richness and frequency of woody angiosperm plants with extrafloral nectaries was studied in the Pasoh Forest Reserve, a primary lowland forest in Peninsular Malaysia. EFN were present on 12.3% of the 741 species surveyed. 91 plant species belonging to 47 genera and 16 families were found to have EFN. Euphorbiaceae, Dipterocarpaceae, Rosaceae, Leguminosae, and Ebenaceae were the families most frequently bearing EFN whereas EFN were rare in the more primitive subclasses of the Magnoliopsida and common in the more advanced taxa (Dilleniidae, Rosidae, Asteridae). Most common were flattened glands associated with the leaf blade. A comparison with data from the Neotropics showed a great similarity in regard to the taxonomic distribution of plants with EFN. EFN-plants occupied 19.3% of the cover of the Pasoh Forest 50 ha plot.Species with EFN were under-represented among shrubs and trees of the understorey. We found an increase in the number of species with EFN as well as in their cover from the understorey to the canopy emergents. EFN were found more often among the abundant species (species with n>500/50 ha). Percentage occurrence and cover of EFN-bearing plants in the 50 ha plot of primary forest was lower than recorded for secondary habitats in tropical areas. At present, in the core zone of the Pasoh Forest Reserve which has been investigated only few species known to indicate disturbance occur. Therefore most of the recorded EFN-species cannot be regarded as secondary forest plants.The interactions between ants and EFN-bearing plants appear to be rather facultative and nonspecific. In Pasoh we found 28 ant species from seven genera visiting EFN. Most of the EFN-associated ants belonged to the subfamily Myrmicinae while Ponerines were rare, a pattern which was also reported for the Neotropics.In cooperation with the Forest Research Institute Malaysia, Kepong, 52109 Kuala Lumpur, Selangor, Malaysia.