Background Termites of the subfamily Macrotermitinae live in a mutualistic symbiosis

Background Termites of the subfamily Macrotermitinae live in a mutualistic symbiosis with basidiomycete fungi of the genus Law and Lewis) only for the entire colony of termites. been found among the symbionts of fungus-growing ants [12,32,33], but here the frequency of sex among symbionts is apparently much lower [34]. Is interaction specificity linked to transmission mode? Symbiont transmission modes have been studied for a rather limited number of macrotermitine species. For the species included in this study, we have direct evidence that Odontotermes badius and Macrotermes natalensis rely on horizontal symbiont acquisition [18,19]; De Fine Licht, Korb and Aanen, pers. obs.). For the other two included species of Odontotermes we have indirect evidence, because: 1. fruiting occurs frequently for both species; 2. all other species of the genus that have so far been tested have horizontal symbiont acquisition [17,20]. Also our inferences for Microtermes have to be based on indirect evidence: fruiting bodies have never been observed for any of BMP2B the species included in the present study and five other species of this genus are known to rely on vertical, uniparental symbiont transmission via the female sex [17,22]. Unexpectedly, the presumably vertically transmitted fungi of the three Microtermes species were shared between species. This indicates that horizontal transmission must Specnuezhenide manufacture occur frequently enough to prevent any host specificity at the species level. In an earlier study [9] it was found that the fungal symbionts of several west African Microtermes species were identical to the symbionts of the divergent genera Ancistrotermes and Synacanthotermes, showing that horizontal transmission also Specnuezhenide manufacture occurred between these genera in this other region. Obviously, the transmission modes for these species need to be addressed directly in future studies. How does interaction specificity arise? For the termite species with horizontal symbiont acquisition, it is still an open question how combinations between termites and their fungal symbionts arise and how specificity arises. One way in which interaction specificity could arise would be geographical isolation, but we did not find strong geographical differentiation at the scale of our study. For some species of fungus-growing termites it has been found that fungal fruiting is synchronized with the time that the first workers of founding colonies of the typical host leave the nest to start foraging [17]. Since dispersal on the wing varies between species, but is highly synchronized within species, such temporal segregation would be a possible way for interaction specificity to arise. Temporal segregation of this kind could potentially explain that all three Odontotermes species in our study were associated with a different range of fungal symbionts, so that none of the ITS haplotypes were shared between the species. A related question of crucial importance is how long basidiospores of Termitomyces remain viable and whether a functional Termitomyces spore Specnuezhenide manufacture bank exists. We hypothesize that Specnuezhenide manufacture active partner selection may also play a role in the establishment of interaction specificity, as neither temporal nor geographical isolation seem sufficient explanations to arrive at the levels of specificity observed. Similarly, for species with predominantly vertical transmission, the question is how symbiont exchange occurs. Detailed experimental studies will be needed to investigate whether and how selection takes place during the establishment and later maintenance of fungal symbionts in termite colonies. Conclusion Interaction specificity between fungus-growing termites and Termitomyces fungi Specnuezhenide manufacture is highly variable and ranges from mutually high specificity to mutually low specificity (Figure ?(Figure1).1). This implies that the hypothesis that inhabitant mutualistic symbionts should be less diverse than the exhabitant hosts is not supported by the.