Florian Menzel

Specialization, constraints, and conflicting interests in mutualistic networks.

The topology of ecological interaction webs holds important information for theories of coevolution, biodiversity, and ecosystem stability . However, most previous network analyses solely counted the number of links and ignored variation in link strength. Because of this crude resolution, results vary with scale and sampling intensity, thus hampering a comparison of network patterns at different levels . We applied a recently developed quantitative and scale-independent analysis based on information theory to 51 mutualistic plant-animal networks, with interaction frequency as measure of link strength. Most networks were highly structured, deviating significantly from random associations. The degree of specialization was independent of network size. Pollination webs were significantly more specialized than seed-dispersal webs, and obligate symbiotic ant-plant mutualisms were more specialized than nectar-mediated facultative ones. Across networks, the average specialization of animal and plants was correlated, but is constrained by the ratio of plant to animal species involved. In pollination webs, rarely visited plants were on average more specialized than frequently attended ones, whereas specialization of pollinators was positively correlated with their interaction frequency. We conclude that quantitative specialization in ecological communities mirrors evolutionary trade-offs and constraints of web architecture. This approach can be easily expanded to other types of biological interactions.

Ecology and Evolution of Communication in Social Insects

Insect life strategies comprise all levels of sociality from solitary to eusocial, in which individuals form persistent groups and divide labor. With increasing social complexity, the need to communicate a greater diversity of messages arose to coordinate division of labor, group cohesion, and concerted actions. Here we summarize the knowledge on prominent messages in social insects that inform about reproduction, group membership, resource locations, and threats and discuss potential evolutionary trajectories of each message in the context of social complexity.

Selective interspecific tolerance in tropical Crematogaster–Camponotus associations

Associations between ants of the genera Crematogaster and Camponotus are found in many parts of the world. Associated species use common trails (trail sharing) or even share a common nest (parabiosis). In a tropical lowland forest in Malaysian Borneo, we studied intraspecific and interspecific aggression among the parabiotic species Crematogaster modiglianii and Camponotus rufifemur using both field and laboratory assays. Cr. modiglianii tolerated Ca. rufifemur workers from certain foreign colonies but fiercely attacked those of others. In contrast, Ca. rufifemur was tolerant even towards attacking allocolonial Cr. modiglianii workers but killed other Crematogaster species. By analogy, other Camponotus species usually attacked and killed Cr. modiglianii. Intraspecific confrontations among Ca. rufifemur colonies yielded a gradient from allocolonial tolerance to strong aggression. The aggression patterns coincide with those of Cr. modiglianii towards Ca. rufifemur workers from the same colonies. Our results suggest either that Ca. rufifemur is not able to recognize allocolonial Cr. modiglianii workers as foreign or that they are recognized but tolerated. The unilateral, species-specific but not colony-specific tolerance of Ca. rufifemur towards its partner species contrasts with highly colony-specific tolerance found among neotropical parabioses.

Tropical parabiotic ants: Highly unusual cuticular substances and low interspecific discrimination

BackgroundAssociations between animal species require that at least one of the species recognizes its partner. Parabioses are associations of two ant species which co-inhabit the same nest. Ants usually possess an elaborate nestmate recognition system, which is based on cuticular hydrocarbons and allows them to distinguish nestmates from non-nestmates through quantitative or qualitative differences in the hydrocarbon composition. Hence, living in a parabiotic association probably necessitates changes of the nestmate recognition system in both species, since heterospecific ants have to be accepted as nestmates.ResultsIn the present study we report highly unusual cuticular profiles in the parabiotic species Crematogaster modiglianii and Camponotus rufifemur from the tropical rainforest of Borneo. The cuticle of both species is covered by a set of steroids, which are highly unusual surface compounds. They also occur in the Dufour gland of Crematogaster modiglianii in high quantities. The composition of these steroids differed between colonies but was highly similar among the two species of a parabiotic nest. In contrast, hydrocarbon composition of Cr. modiglianii and Ca. rufifemur differed strongly and only overlapped in three regularly occurring and three trace compounds. The hydrocarbon profile of Camponotus rufifemur consisted almost exclusively of methyl-branched alkenes of unusually high chain lengths (up to C49). This species occurred in two sympatric, chemically distinct varieties with almost no hydrocarbons in common. Cr. modiglianii discriminated between these two varieties. It only tolerated workers of the Ca. rufifemur variety it was associated with, but attacked the respective others. However, Cr. modiglianii did not distinguish its own Ca. rufifemur partner from allocolonial Ca. rufifemur workers of the same variety.ConclusionWe conclude that there is a mutual substance transfer between Cr. modiglianii and Ca. rufifemur. Ca. rufifemur actively or passively acquires cuticular steroids from its Cr. modiglianii partner, while the latter acquires at least two cuticular hydrocarbons from Ca. rufifemur. The cuticular substances of both species are highly unusual regarding both substance classes and chain lengths, which may cause the apparent inability of Cr. modiglianii to discriminate Ca. rufifemur nestmates from allocolonial Ca. rufifemur workers of the same chemical variety.

Worker personality and its association with spatially structured division of labor.

Division of labor is a defining characteristic of social insects and fundamental to their ecological success. Many of the numerous tasks essential for the survival of the colony must be performed at a specific location. Consequently, spatial organization is an integral aspect of division of labor. The mechanisms organizing the spatial distribution of workers, separating inside and outside workers without central control, is an essential, but so far neglected aspect of division of labor. In this study, we investigate the behavioral mechanisms governing the spatial distribution of individual workers and its physiological underpinning in the ant Myrmica rubra. By investigating worker personalities we uncover position-associated behavioral syndromes. This context-independent and temporally stable set of correlated behaviors (positive association between movements and attraction towards light) could promote the basic separation between inside (brood tenders) and outside workers (foragers). These position-associated behavior syndromes are coupled with a high probability to perform tasks, located at the defined position, and a characteristic cuticular hydrocarbon profile. We discuss the potentially physiological causes for the observed behavioral syndromes and highlight how the study of animal personalities can provide new insights for the study of division of labor and self-organized processes in general.

TOLERANCE REQUIRES THE RIGHT SMELL: FIRST EVIDENCE FOR INTERSPECIFIC SELECTION ON CHEMICAL RECOGNITION CUES

The integument of insects is generally covered with cuticular hydrocarbons (CHC). They serve multiple functions, most prominent among them waterproofing and—especially among social insects—as communication signal. CHC profiles are incredibly diverse within and across species. However, the causes for CHC variation between species, and potential selection pressures that may shape CHC profiles, are hardly understood. Here, we investigated potential selection pressures on ant CHC. We tested the hypotheses that living in association with another species (e.g., parabiosis), and the climate of the ants habitat, affect CHC composition. We conducted a large‐scale comparison of 37 Camponotus species from five continents. Our results demonstrate that closely associated ant species possess significantly longer hydrocarbons and higher proportions of methylbranched alkenes and alkadienes than non‐ or loosely associated species. In contrast, climatic factors had no effects. This study shows that the need to be tolerated by another species greatly affects CHC profiles.

Trail-sharing among tropical ants: interspecific use of trail pheromones?

1. Trail‐sharing between different ant species is rare and restricted to a small number of species pairs. Its underlying mechanisms are largely unknown. For trail‐sharing to occur, two factors are required: (i) one or both species must recognise the other species or its pheromone trails and (ii) both species must tolerate each other to a certain extent to allow joint use of the trail. A species that follows anothers trails can efficiently exploit the others information on food sources contained in the pheromone trails. Hence, food competition and thus aggressive interactions between a species following anothers trail and the species being followed, seem likely.

Behavioural and chemical mechanisms behind a Mediterranean ant-ant association

1. Interspecific competition among ants is common, and so is competitive exclusion among dominant ant species. In contrast, specific associations between non‐parasitic ant species are rare, especially in the temperate zones. As an exception, the subordinate ant Camponotus lateralis frequently co‐occurs with the dominant Crematogaster scutellaris but rarely with other dominant ants.

Covariation and phenotypic integration in chemical communication displays: biosynthetic constraints and eco-evolutionary implications

Chemical communication is ubiquitous. The identification of conserved structural elements in visual and acoustic communication is well established, but comparable information on chemical communication displays (CCDs) is lacking. We assessed the phenotypic integration of CCDs in a meta-analysis to characterize patterns of covariation in CCDs and identified functional or biosynthetically constrained modules. Poorly integrated plant CCDs (i.e. low covariation between scent compounds) support the notion that plants often utilize one or few key compounds to repel antagonists or to attract pollinators and enemies of herbivores. Animal CCDs (mostly insect pheromones) were usually more integrated than those of plants (i.e. stronger covariation), suggesting that animals communicate via fixed proportions among compounds. Both plant and animal CCDs were composed of modules, which are groups of strongly covarying compounds. Biosynthetic similarity of compounds revealed biosynthetic constraints in the covariation patterns of plant CCDs. We provide a novel perspective on chemical communication and a basis for future investigations on structural properties of CCDs. This will facilitate identifying modules and biosynthetic constraints that may affect the outcome of selection and thus provide a predictive framework for evolutionary trajectories of CCDs in plants and animals.

Intraspecific nestmate recognition in two parabiotic ant species: acquired recognition cues and low inter-colony discrimination

Parabiotic ants—ants that share their nest with another ant species—need to tolerate not only conspecific nestmates, but also nestmates of a foreign species. The parabiotic ants Camponotus rufifemur and Crematogaster modiglianii display high interspecific tolerance, which exceeds their respective partner colony and extends to alien colonies of the partner species. The tolerance appears to be related to unusual cuticular substances in both species. Both species possess hydrocarbons of unusually high chain lengths. In addition, Cr. modiglianii carries high quantities of hereto unknown compounds on its cuticle. These unusual features of the cuticular profiles may affect nestmate recognition within both respective species as well. In the present study, we therefore examined inter-colony discrimination within the two parabiotic species in relation to chemical differentiation. Cr. modiglianii was highly aggressive against workers from alien conspecific colonies in experimental confrontations. In spite of high inter-colony variation in the unknown compounds, however, Cr. modiglianii failed to differentiate between intracolonial and allocolonial unknown compounds. Instead, the cuticular hydrocarbons functioned as recognition cues despite low variation across colonies. Moreover, inter-colony aggression within Cr. modiglianii was significantly influenced by the presence of two methylbranched alkenes acquired from its Ca. rufifemur partner. Ca. rufifemur occurs in two varieties (‘red’ and ‘black’) with almost no overlap in their cuticular hydrocarbons. Workers of this species showed low aggression against conspecifics from foreign colonies of the same variety, but attacked workers from the respective other variety. The low inter-colony discrimination within a variety may be related to low chemical differentiation between the colonies. Ca. rufifemur majors elicited significantly more inter-colony aggression than medium-sized workers. This may be explained by the density of recognition cues: majors carried significantly higher quantities of cuticular hydrocarbons per body surface.