Adrian Brückner

Surface area–volume ratios in insects

Body mass, volume and surface area are important for many aspects of the physiology and performance of species. Whereas body mass scaling received a lot of attention in the literature, surface areas of animals have not been measured explicitly in this context. We quantified surface area–volume (SA/V) ratios for the first time using 3D surface models based on a structured light scanning method for 126 species of pollinating insects from 4 orders (Diptera, Hymenoptera, Lepidoptera, and Coleoptera). Water loss of 67 species was measured gravimetrically at very dry conditions for 2 h at 15 and 30 °C to demonstrate the applicability of the new 3D surface measurements and relevance for predicting the performance of insects. Quantified SA/V ratios significantly explained the variation in water loss across species, both directly or after accounting for isometric scaling (residuals of the SA/V ∼ mass2/3 relationship). Small insects with a proportionally larger surface area had the highest water loss rates. Surface scans of insects to quantify allometric SA/V ratios thus provide a promising method to predict physiological responses, improving the potential of body mass isometry alone that assume geometric similarity.

Storage and release of hydrogen cyanide in a chelicerate (Oribatula tibialis)

Significance Hydrogen cyanide (HCN) is highly volatile and among the most toxic substances known, being lethal to humans at a dosage of 1–2 mg/kg body weight. HCN blocks the respiratory chain and prevents aerobic organisms from using oxygen. In nature, HCN is produced by numerous plants that store it mainly as glycosides. Among animals, cyanogenesis is a defensive strategy that has seemed restricted to a few mandibulate arthropods (certain insects, millipedes, and centipedes), which evolved ways to store HCN in the form of stable and less volatile molecules. We found an instance of cyanogenesis in the phylogenetically distant group Chelicerata (“spider-like” arthropods), involving an aromatic ester for stable HCN storage and two degradation pathways that release HCN. Cyanogenesis denotes a chemical defensive strategy where hydrogen cyanide (HCN, hydrocyanic or prussic acid) is produced, stored, and released toward an attacking enemy. The high toxicity and volatility of HCN requires both chemical stabilization for storage and prevention of accidental self-poisoning. The few known cyanogenic animals are exclusively mandibulate arthropods (certain myriapods and insects) that store HCN as cyanogenic glycosides, lipids, or cyanohydrins. Here, we show that cyanogenesis has also evolved in the speciose Chelicerata. The oribatid mite Oribatula tibialis uses the cyanogenic aromatic ester mandelonitrile hexanoate (MNH) for HCN storage, which degrades via two different pathways, both of which release HCN. MNH is emitted from exocrine opisthonotal oil glands, which are potent organs for chemical defense in most oribatid mites.

Scent of a mite: origin and chemical characterization of the lemon-like flavor of mite-ripened cheeses.

Cheese infested with cheese mites is usually treated as unpalatable. Nevertheless, some traditional cheese manufactories in Germany and France intentionally use mites for fermentation of special varieties (i.e. Milbenkäse and Mimolette). While their production includes different mite species, both are characterized by a “lemon-like” flavor. However, the chemical nature and origin of this flavor-component is unknown. The cheese mites possess a pair of opisthosomal glands producing blends of hydrocarbons, terpenes and aromatics. Here, we describe the chemical profiles of the astigmatid mite species Tyrolichus casei (Milbenkäse) and Acarus siro (Mimolette). Although the chemical profiles differ in several aspects, both mite species produce neral (a volatile flavor component of lemon oil), which was absent from the headspace of both cheeses without mites. We conclude that the lemon-like flavor of mite cheese is not a consequence of fermentation of the cheese itself but a component from secretions of the cheese mites.

Nutrient quality of vertebrate dung as a diet for dung beetles

At the basis of a trophic web, coprophagous animals like dung beetles (Scarabaeoidea) utilize resources that may have advantages (easy gain and handling) as well as drawbacks (formerly processed food). Several studies have characterized the nutrients, e.g. C/N ratios and organic matter content, for specific types of dung. However, a comparative approach across dung types and feeding guilds of dung producers, and relationships between dung nutrients and preferences by coprophages, have been missing. Hence, we analyzed water content, C/N ratio, amino acid, neutral lipid fatty acid, free fatty acid and sterol composition and concentrations in dung from 23 vertebrates, including carnivore, omnivore and herbivore species. Our analyses revealed significant differences among the three vertebrate feeding guilds for most nutritional parameters. Although formerly processed, dung grants sufficient amounts of essential nutrients for insects. We tested whether nutrients can explain the dung beetles’ preferences in a field experiment, using 12 representative dung types in baits that were installed in 27 forests and 27 grasslands. Although consistent preferences for specific dung types were pronounced, the nutritional composition did not predict the variation in attractiveness of these dung diets, suggesting a primary role of dung volatiles irrespective of food quality.

The relationship between epicuticular long-chained hydrocarbons and surface area - volume ratios in insects (Diptera, Hymenoptera, Lepidoptera)

Long-chain cuticular hydrocarbons (CHCs) are common components of the epicuticle of terrestrial arthropods. CHC serve as a protective barrier against environmental influences but also act as semiochemicals in animal communication. Regarding the latter aspect, species- or intra-functional group specific CHCs composition and variation are relatively well studied. However, comparative knowledge about the relationship of CHC quantity and their relation to surface area—volume ratios in the context of water loss and protection is fragmentary. Hence, we aim to study the taxon-specific relationship of the CHC amount and surface-area to volume ratio related to their functional role (e.g. in water loss). We focused on flower visiting insects and analyzed the CHC amounts of three insect orders (Hymenoptera, Lepidoptera and Diptera) using gas chromatography—mass spectrometry (GC-MS). We included 113 species from two grassland plots, quantified their CHCs, and measured their body mass and surface area. We found differences in the surface area, CHCs per body mass and the CHC density (= amount of CHCs per surface area) across the three insect taxa. Especially the Hymenoptera had a higher CHC density compared to Diptera and Lepidoptera. CHC density could be explained by surface area-volume ratios in Hymenoptera but not in Diptera and Lepidoptera. Unexpectedly, CHC density decreased with increasing surface area—volume ratios.

Patterns and dynamics of neutral lipid fatty acids in ants – implications for ecological studies

BackgroundTrophic interactions are a fundamental aspect of ecosystem functioning, but often difficult to observe directly. Several indirect techniques, such as fatty acid analysis, were developed to assess these interactions. Fatty acid profiles may indicate dietary differences, while individual fatty acids can be used as biomarkers. Ants are among the most important terrestrial animal groups, but little is known about their lipid metabolism, and no study so far used fatty acids to study their trophic ecology. We set up a feeding experiment with high- and low-fat food to elucidate patterns and dynamics of neutral lipid fatty acids (NLFAs) assimilation in ants. We asked whether dietary fatty acids are assimilated through direct trophic transfer, how diet influences NLFA total amounts and patterns over time, and whether these assimilation processes are similar across species and life stages.ResultsAnts fed with high-fat food quickly accumulated specific dietary fatty acids (C18:2n6, C18:3n3 and C18:3n6), compared to ants fed with low-fat food. Dietary fat content did not affect total body fat of workers or amounts of fatty acids extensively biosynthesized by animals (C16:0, C18:0, C18:1n9). Larval development had a strong effect on the composition and amounts of C16:0, C18:0 and C18:1n9. NLFA compositions reflected dietary differences, which became more pronounced over time. Assimilation of specific dietary NLFAs was similar regardless of species or life stage, but these factors affected dynamics of other NLFAs, composition and total fat.ConclusionsWe showed that ants accumulated certain dietary fatty acids via direct trophic transfer. Fat content of the diet had no effect on lipids stored by ants, which were able to synthesize high amounts of NLFAs from a sugar-based diet. Nevertheless, dietary NLFAs had a strong effect on metabolic dynamics and profiles. Fatty acids are a useful tool to study trophic biology of ants, and could be applied in an ecological context, although factors that affect NLFA patterns should be taken into account. Further studies should address which NLFAs can be used as biomarkers in natural ant communities, and how factors other than diet affect fatty acid dynamics and composition of species with distinct life histories.

Chemical and behavioral integration of army ant-associated rove beetles - a comparison between specialists and generalists

Host-symbiont interactions are embedded in ecological communities and range from unspecific to highly specific relationships. Army ants and their arthropod guests represent a fascinating example of species-rich host-symbiont associations where host specificity ranges across the entire generalist - specialist continuum. In the present study, we compared the behavioral and chemical integration mechanisms of two extremes of the generalist - specialist continuum: generalist ant-predators in the genus Tetradonia (Staphylinidae: Aleocharinae: Athetini), and specialist ant-mimics in the genera Ecitomorpha and Ecitophya (Staphylinidae: Aleocharinae: Ecitocharini). Similar to a previous study of Tetradonia beetles, we combined DNA barcoding with morphological studies to define species boundaries in ant-mimicking beetles. This approach found four ant-mimicking species at our study site at La Selva Biological Station in Costa Rica. Community sampling of Eciton army ant parasites revealed that ant-mimicking beetles were perfect host specialists, each beetle species being associated with a single Eciton species. These specialists were seamlessly integrated into the host colony, while generalists avoided physical contact to host ants in behavioral assays. Analysis of the ants’ nestmate recognition cues, i.e. cuticular hydrocarbons (CHCs), showed close similarity in CHC composition and CHC concentration between specialists and Eciton burchellii foreli host ants. On the contrary, the chemical profiles of generalists matched host profiles less well, indicating that high accuracy in chemical host resemblance is only accomplished by socially integrated species. Considering the interplay between behavior, morphology, and cuticular chemistry, specialists but not generalists have cracked the ants’ social code with respect to various sensory modalities. Our results support the long-standing idea that the evolution of host-specialization in parasites is a trade-off between the range of potential host species and the level of specialization on any particular host.

The ontogeny of oil gland chemistry in the oribatid mite Archegozetes longisetosus Aoki (Oribatida, Trhypochthoniidae)

ABSTRACT The so-called “glandulate Oribatida” (including Astigmata) possess large opisthonotal oil glands, which produce semiochemicals as potent agents for chemical defence against predators (allomones) as well as for intraspecific communication (pheromones). The oil gland reservoirs and their contents remain with shed exuviae, so secretions need to be synthesized de novo after each moult. The qualitative composition of chemical profiles may change dramatically between the last juvenile instar (tritonymph) and adult (e.g. Hermannia convexa Koch, Scheloribates azumaensis Enami, Nakamura & Katsumata, Oribotritia berlesei Michael), but reports about quantitative ontogenetic shifts are ambiguous. Here, we analysed the complete ontogenetic sequence (larva, protonymph, deutonymph, tritonymph, and adult) of oil gland secretions in the parthenogenetic oribatid model species Archegozetes longisetosus Aoki using gas chromatography–mass spectrometry. We show that absolute and body mass-corrected amounts of oil gland secretions increase during ontogeny and that secretion quantity scales allometrically with body mass (i.e. ontogenetic instar). Furthermore, we found highly significant ontogenetic shifts in the relative quantity of chemical components among instars, but the qualitative composition (=2,6-HMBD, neral, neryl formate, γ-acaridial, tridecane, 7-pentadecene, pentadecane, 6,9-heptadecadiene, 8-heptadecene, and heptadecane) remained stable.

Seasonal fluctuation of oribatid mite communities in forest microhabitats

Oribatid mites are abundant and diverse decomposers in almost all terrestrial microhabitats, especially in temperate forests. Although their functional importance in the decomposition system in these forests has been investigated, spatio-temporal patterns of oribatid mite communities inhabiting different microhabitats have largely been neglected. Therefore, we (i) investigated seasonal fluctuation (monthly over one year) in oribatid-mite community structure and specificity to three microhabitats (moss, dead wood and litter) and (ii) analyzed the influence of air temperature and overall air humidity on seasonal community changes. In total, 57,398 adult oribatid mite individuals were collected. Total abundance, species richness and diversity differed among microhabitats. Seasonal changes were most pronounced in moss and least in litter. While overall air humidity had no influence on species distribution and community changes, air temperature positively influenced species richness and diversity, again most pronounced in moss. The calculated environmental temperature occurrence niche showed that 35% of adult oribatid mite species occurred at higher air temperatures. Furthermore, interaction/bipartite networks were more generalized—i.e., species were more equally distributed among moss, dead wood and litter—when ambient air temperatures were higher. This pattern is probably due to the dispersal ability of adult oribatid mites, i.e., species enter a dispersal mode only at higher air temperatures.

Unveiling community patterns and trophic niches of tropical and temperate ants using an integrative framework of field data, stable isotopes and fatty acids

Background The use and partitioning of trophic resources is a central aspect of community function. On the ground of tropical forests, dozens of ant species may be found together and ecological mechanisms should act to allow such coexistence. One hypothesis states that niche specialization is higher in the tropics, compared to temperate regions. However, trophic niches of most species are virtually unknown. Several techniques might be combined to study trophic niche, such as field observations, fatty acid analysis (FAA) and stable isotope analysis (SIA). In this work, we combine these three techniques to unveil partitioning of trophic resources in a tropical and a temperate community. We describe patterns of resource use, compare them between communities, and test correlation and complementarity of methods to unveil both community patterns and species’ niches. Methods Resource use was assessed with seven kinds of bait representing natural resources available to ants. Neutral lipid fatty acid (NLFA) profiles, and δ15N and δ13C isotope signatures of the species were also obtained. Community patterns and comparisons were analyzed with clustering, correlations, multivariate analyses and interaction networks. Results Resource use structure was similar in both communities. Niche breadths (H′) and network metrics (Q and H2′) indicated similar levels of generalization between communities. A few species presented more specialized niches, such as Wasmannia auropunctata and Lasius fuliginosus. Stable isotope signatures and NLFA profiles also indicated high generalization, although the latter differed between communities, with temperate species having higher amounts of fat and proportions of C18:1n9. Bait use and NLFA profile similarities were correlated, as well as species’ specialization indices (d′) for the two methods. Similarities in δ15N and bait use, and in δ13C and NLFA profiles, were also correlated. Discussion Our results agree with the recent view that specialization levels do not change with latitude or species richness. Partition of trophic resources alone does not explain species coexistence in these communities, and might act together with behavioral and environmental mechanisms. Temperate species presented NLFA patterns distinct from tropical ones, which may be related to environmental factors. All methods corresponded in their characterization of species’ niches to some extent, and were robust enough to detect differences even in highly generalized communities. However, their combination provides a more comprehensive picture of resource use, and it is particularly important to understand individual niches of species. FAA was applied here for the first time in ant ecology, and proved to be a valuable tool due to its combination of specificity and temporal representativeness. We propose that a framework combining field observations with chemical analysis is valuable to understand resource use in ant communities.