Many plants interact with carnivores as an indirect defence against herbivores. The release of volatile organic compounds (VOCs) and the secretion of extrafloral nectar (EFN) are induced by insect feeding, a response that is mediated by the plant hormone, jasmonic acid. Although VOCs mainly attract predatory mites and parasitic wasps, while EFN mainly attracts ants, many more animal-plant interactions are influenced by these two traits. Other traits involved in defensive tritrophic interactions are cellular food bodies and domatia, which serve the nutrition and housing of predators. They are not known to respond to herbivory, while food body production can be induced by the presence of the mutualists. Interactions among the different defensive traits, and between them and other biotic and abiotic factors exist on the genetic, physiological, and ecological levels, but so far remain understudied. Indirect defences are increasingly being discussed as an environmentally-friendly crop protection strategy, but much more knowledge on their fitness effects under certain environmental conditions is required before we can understand their ecological and evolutionary relevance, and before tritrophic interactions can serve as a reliable tool in agronomy.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2007.02330.x

Indirect defence via tritrophic interactions.

▪ Abstract Protective ant-plant interactions, important in both temperate and tropical communities, are increasingly used to study a wide range of phenomena of general interest. As antiherbivore defenses “worn on the outside,” they pose fewer barriers to experimentation than do direct (e.g., chemical) plant defenses. This makes them tractable models to study resource allocation to defense and mechanisms regulating it. As multi-trophic level interactions varying in species specificity and impact on fitness of participants, ant-plant-herbivore associations figure prominently in studies of food-web structure and functioning. As horizontally transmitted mutualisms that are vulnerable to parasites and “cheaters,” ant-plant symbioses are studied to probe the evolutionary dynamics of interspecies interactions. These symbioses, products of coevolution between plants and insect societies, offer rich material for studying ant social evolution in novel contexts, in settings where colony limits, resource supply, and ...

Protective ant-plant interactions as model systems in ecological and evolutionary research.

A catalog of the cecidomyiidae (Diptera) of the world

Interactions between ants and honeydew-producing hemipteran insects are abundant and widespread in arthropod food webs, yet their ecological consequences are very poorly known. Ant–hemipteran interactions have potentially broad ecological effects, because the presence of honeydew-producing hemipterans dramatically alters the abundance and predatory behaviour of ants on plants. We review several studies that investigate the consequences of ant–hemipteran interactions as ‘keystone interactions’ on arthropod communities and their host plants. Ant–hemipteran interactions have mostly negative effects on the local abundance and species richness of several guilds of herbivores and predators. In contrast, out of the 30 studies that document the effects of ant–hemipteran interactions on plants, the majority (73%) shows that plants actually benefit indirectly from these interactions. In these studies, increased predation or harassment of other, more damaging, herbivores by hemipteran-tending ants resulted in decreased plant damage and/or increased plant growth and reproduction. The ecological consequences of mutualistic interactions between honeydew-producing hemipterans and invasive ants relative to native ants have rarely been studied, but they may be of particular importance owing to the greater abundance, aggressiveness and extreme omnivory of invasive ants. We argue that ant–hemipteran interactions are largely overlooked and underappreciated interspecific interactions that have strong and pervasive effects on the communities in which they are embedded.

https://royalsocietypublishing.org/doi/pdf/10.1098/rspb.2006.3701

Ecological consequences of interactions between ants and honeydew-producing insects.

Specialization, constraints, and conflicting interests in mutualistic networks.

Ant-plant interactions in the canopy of a lowland Amazonian rainforest of the upper Orinoco, Venezuela, were studied using a modified commercial crane on rails (Surumoni project). Our observations show a strong correlation between plant sap exudates and both abundance of ants and co-occurrence of ant species in tree canopies. Two types of plant sap sources were compared: extrafloral nectaries (EFNs) and honeydew secretions by homopterans. EFNs were a frequent food source for ants on epiphytes (Philodendron spp., Araceae) and lianas (Dioclea, Fabaceae), but rare on canopy trees in the study area, whereas the majority of trees were host to aggregations of homopterans tended by honeydew-seeking ants (on 62% of the trees examined). These aggregations rarely occurred on epiphytes. Baited ant traps were installed on plants with EFNs and in the crowns of trees from three common genera, including trees with and without ant-tended homopterans: Goupia glabra (Celastraceae), Vochysia spp. (Vochysiaceae), and Xylopia spp. (Annonaceae). The number of ant workers per trap was significantly higher on plants offering one of the two plant sap sources than on trees without such resources. Extrafloral nectaries were used by a much broader spectrum of ant species and genera than honeydew, and co-occurrence of ant species (in traps) was significantly higher on plants bearing EFNs than on trees. Homopteran honeydew (Coccidae and Membracidae), on the other hand, was mostly monopolised by a single ant colony per tree. Homopteran-tending ants were generally among the most dominant ants in the canopy. The most prominent genera were Azteca, Dolichoderus (both Dolichoderinae), Cephalotes, Pheidole, Crematogaster (all Myrmicinae), and Ectatomma (Ponerinae). Potential preferences were recorded between ant and homopteran species, and also between ant-homopteran associations and tree genera. We hypothesize that the high availability of homopteran honeydew provides a key resource for ant mosaics, where dominant ant colonies and species maintain mutually exclusive territories on trees. In turn, we propose that for nourishment of numerous ants of lower competitive capacity, Philodendron and other sources of EFNs might be particularly important.

How plants shape the ant community in the Amazonian rainforest canopy: the key role of extrafloral nectaries and homopteran honeydew.

In 2005, researchers at the Leipzig Canopy Crane Research Facility collected living leaves of four temperate tree species at heights of between 15 and 33 m above the ground. Following surface sterilisation of the leaves, leaf-fragments were cultured on malt extract agar which allowed the growth of endophytic fungi into the surrounding medium. Isolated cultures were identified by morphology and sequence analysis of the D1/D2 region of the large subunit rDNA. Phylogenetic analysis established the taxonomic positions of the fungi. A total of 49 different taxa were identified, representing 20 families and ten orders. With the exception of one basidiomycetous yeast, all taxa belonged to filamentous ascomycetes. Species richness was highest on Tilia cordata and lowest on Quercus robur. Species-accumulation curves showed that the sampling effort was not sufficient to cover the majority of the likely species at the investigation site. Most endophytes proved to be ubiquitous within the canopy of the investigation site, but habitat preferences in terms of different tree species, different light regimes and season (sampling times) were obvious for some abundant endophytes. Apiognomonia errabunda and Aspergillus niger occurred predominantly on Q. robur, Diplodina acerina on Acer pseudoplatanus, one species of Phoma significantly prefered shaded leaves from the lower canopy layer whereas Sordaria fimicola prefered sun-exposed leaves from the upper tree crowns. Seasonal patterns were observed, for example, for A. errabunda, which was abundant in young leaves in the spring and almost completely absent in aged autumn-leaves, thus suggesting the accumulation of antifungal secondary plant metabolites during the growing season.

Species richness and distribution patterns of leaf-inhabiting endophytic fungi in a temperate forest canopy

Summary Cornaceae, and Hydrangeaceae point to a very close relationship between the Viburnaceae, Sambucaceae and Adoxaceae, corroborating the inclusion of these families into an order different from the Dipsacales, namely the Viburnales. Many cytological features shared by these families differ strongly from the Dipsacales s.str., especially (1) chromosome size and morphology, (2) the presence of cold induced chromosome regions (CIRs), (3) interphase nuclear structure, and (4) chromosome condensing behaviour at prophase. Cornaceae and Hydrangeaceae present similar interphase nuclei, but differ from the Viburnales by other karyomorphological characters. The results are discussed with respect to previous morphological, embryological, and molecular findings.

Viburnales: cytological features and a new circumscription

In the more than twenty years in which long-term canopy research has been conducted, mycology has been largely disregarded. Our studies using a construction crane to gain access to the canopy of a forest in Leipzig, Germany are the first long term investigations assessing the diversity and ecology of wood-decaying fungi in a canopy. Thirty-seven individuals of nine different tree species with a large amount of dead wood were selected. Sampling focussed on the four most prominent tree species Acer pseudoplatanus, Fraxinus excelsior, Quercus robur and Tilia cordata. In the years 2002 and 2003 dead wood was collected in different canopy strata. Dead branches were removed and stored for two weeks in open boxes with high humidity to allow growth of fructifications in the laboratory. 118 different taxa were identified (108 species, 77 genera). Corticioid fungi (e.g., of Corticiaceae, Stereaceae, Hymenochaetaceae) dominated the fungal composition with 37 species, pyrenomycetes were present with 18 species. Agaric fungi (Agaricales and Cortinariales) were scarce. Species with minute basidiomes dominated the fungal composition of this systematic group. Agarics with larger sporomes were found only once and were restricted to strongly decayed branches in shaded canopy areas. Concerning species richness and fungal composition the four tree species mentioned above differed remarkably. As expected, many fungi that grew on bark or slightly decayed wood showed a distinct host and substratum specifity. It is noteworthy that fungi which are purportedly to be non-specific were found on single tree species only.

/pdf/species-richness-and-substrate-specificity-of-lignicolous-1y9d3rq23h.pdf

Species richness and substrate specificity of lignicolous fungi in the canopy of a temperate, mixed deciduous forest

The genera Viburnum, Sambucus and Lonicera have been investigated for chromosome number and karyomorphology including Giemsa-C-banding, fluorochrome (DAPI/CMA) banding and cold treatment. Cold-induced undercontracted chromosome regions (CIRs) are found in Viburnum and Sambucus for the first time and are apparently identical with larger hc regions, shown by Giemsa C-banding. Certain narrow C-bands are not cold-sensitive. CIRs frequently react brightly CMA-positive in Viburnum and Sambucus, while DAPI fluorescence is virtually ineffective. The occurrence of CIRs within plants is possibly linked to certain nuclear characters such as large chromosomes and continuous condensation behaviour. Cold-induction has possibly also some influence on euchromatin condensation characteristics in prophasic chromosomes.



Several karyological characters point to a closer relationship between Viburnum, Sambucus and Adoxa: Relatively large chromosomes, continuous condensation behaviour, reticulate to semireticulate interphase nuclei and presence of CIRs. These genera appear isolated from Lonicera and the Caprifoliaceae s.str., which differ remarkably in karyomorphology.

Wilfried Morawetz

Papers

How plants shape the ant community in the Amazonian rainforest canopy: the key role of extrafloral nectaries and homopteran honeydew.

Species richness and distribution patterns of leaf-inhabiting endophytic fungi in a temperate forest canopy

Viburnales: cytological features and a new circumscription

Species richness and substrate specificity of lignicolous fungi in the canopy of a temperate, mixed deciduous forest

Cold‐induced Chromosome Regions and Karyosystematics in Sambucus and Viburnum