Biodiversity is important for ecosystem functioning and biotic interactions. In experimental grasslands, increasing plant species richness is known to increase the diversity of associated herbivores and their predators. If these interactions can also involve endosymbionts that reside within a plant or animal host is currently unknown. In plant-feeding aphids, secondary bacterial symbionts can have strong fitness effects on the host, e.g. resistance to natural enemies or fungal pathogens. We examined the secondary symbiont community in three species of aphid, each feeding on a unique host plant across experimental plots that varied in plant species richness. Aphids were collected in May and June, and the symbiont community identified using species-specific PCR assays. Aphis fabae aphids were found to host six different symbiont species with individual aphids co-hosting up to four symbionts. Uroleucon jaceae and Macrosiphum rosae hosted two and three symbiont species, respectively. We found that, at the aphid population level, increasing plant species richness increased the diversity of the aphid symbiont community, whereas at the individual aphid level, the opposite was found. These effects are potentially driven by varying selective pressures across different plant communities of varying diversities, mediated by defensive protection responses and a changing cost-benefit trade-off to the aphid for hosting multiple secondary symbionts. Our work extends documented effects of plant diversity beyond visible biotic interactions to changes in endosymbiont communities, with potentially far-reaching consequences to related ecosystem processes.
Plants exhibit impressive genetic and chemical diversity, not just between species but also within species, and the importance of plant intraspecific variation for structuring ecological communities is well known. When there is variation at the local population level, this can create a spatially heterogeneous habitat for specialised herbivores potentially leading to non‐random distribution of individuals across host plants. Plant variation can affect herbivores directly and indirectly via a third species, resulting in variable herbivore growth rates across different host plants. Herbivores also exhibit within‐species variation, with some genotypes better adapted to some plant variants than others. We genotyped aphids collected across 2 years from a field site containing ~200 patchily distributed host plants that exhibit high chemical diversity. The distribution of aphid genotypes, their ant mutualists, and other predators was assessed across the plants. We present evidence that the local distribution of aphid (Metopeurum fuscoviride) genotypes across host‐plant individuals is associated with variation in the plant volatiles (chemotypes) and non‐volatile metabolites (metabotypes) of their host plant tansy (Tanacetum vulgare). Furthermore, these interactions in the field were influenced by plant‐host preferences of aphid‐mutualist ants. Our results emphasise that plant intraspecific variation can structure ecological communities not only at the species level but also at the genetic level within species and that this effect can be enhanced through indirect interactions with a third species.
There is a great demand for standardising biodiversity assessments in order to allow optimal comparison across research groups. For invertebrates, pitfall or flight-interception traps are commonly used, but sampling solution differs widely between studies, which could influence the communities collected and affect sample processing (morphological or genetic). We assessed arthropod communities with flight-interception traps using three commonly used sampling solutions across two forest types and two vertical strata. We first considered the effect of sampling solution and its interaction with forest type, vertical stratum, and position of sampling jar at the trap on sample condition and community composition. We found that samples collected in copper sulphate were more mouldy and fragmented relative to other solutions which might impair morphological identification, but condition depended on forest type, trap type and the position of the jar. Community composition, based on order-level identification, did not differ across sampling solutions and only varied with forest type and vertical stratum. Species richness and species-level community composition, however, differed greatly among sampling solutions. Renner solution was highly attractant for beetles and repellent for true bugs. Secondly, we tested whether sampling solution affects subsequent molecular analyses and found that DNA barcoding success was species-specific. Samples from copper sulphate produced the fewest successful DNA sequences for genetic identification, and since DNA yield or quality was not particularly reduced in these samples additional interactions between the solution and DNA must also be occurring. Our results show that the choice of sampling solution should be an important consideration in biodiversity studies. Due to the potential bias towards or against certain species by Ethanol-containing sampling solution we suggest ethylene glycol as a suitable sampling solution when genetic analysis tools are to be used and copper sulphate when focusing on morphological species identification and facing financial restrictions in biodiversity studies.
Habitat restoration aims to improve local habitat conditions for threatened species. While such restorations are widespread, rigorous evaluations of their success are rare. This is especially true of those considering species dynamics. Increasingly, deadwood is a target for forest restoration as many species directly and indirectly depend on this resource. In a broadleaf forest in southern Germany, we explored the effect of landscape‐wide deadwood restoration on the population genetic structure of the specialist fungus‐dwelling saproxylic beetle Bolitophagus reticulatus. Before 2003, the northern area of this forest was intensively logged for more than half a century, while the southern part was less intensively managed. This drove populations of the host fungus Fomes fomentarius, and consequently the beetle, to local extinction in the northern part. Only after the first decade of restoration were both the fungus and its beetles present across all areas of the forest. Using 17 newly developed microsatellite loci, we show that these beetles exhibit population genetic structuring, mainly influenced by the north‐south divide. However, the low degree of isolation‐by‐distance, and the low relatedness of beetles collected from the same trees or fungus, shows that strong dispersal ability is facilitating the recolonisation of these forests on the scale of tens of kilometers. In another 10 years, it is likely that the population will show even less genetic structuring. Synthesis and applications. Through the recolonisation of the fungus Fomes fomentarius and the fungus‐dwelling beetle Bolitophagus reticulatus after deadwood restoration, we demonstrate that, while there are many discussions on the optimal spatial distribution of deadwood, just the presence of deadwood can be sufficient to enable recolonisations of specific species. As long as some relict populations of these species are embedded in a once intensively managed forest, increased deadwood (amount and diversity) anywhere will benefit recolonisation of the habitat. However, increasing deadwood diversity should also be encouraged to benefit even more species.
1. Floral plantings adjacent to crops fields can recruit populations of natural enemies by providing flower nectar and non-crop prey to increase natural pest regulation. Observed variation in success rates might be due to changes in the unseen community of protective endosymbionts hosted by many herbivorous insects, which can confer resistance to various specialist natural enemies, e.g. parasitoid wasps. Reduced insect control may occur if highly protective symbiont combinations increase in frequency via selection effects, and this is expected to be stronger in lower diversity systems. 2. We used a large-scale field trial to analyse the bacterial endosymbiont communities hosted by cereal aphids (Sitobion avenae) collected along transects into strip plots of barley plants managed by either conventional or integrated (including floral field margins and reduced inputs) methods. In addition, we conducted an outdoor pot experiment to analyse endosymbionts in S. avenae aphids collected on barley plants that were either grown alone or alongside one of three flowering plants, across three time points. 3. In the field, aphids hosted up to four symbionts. The abundance of aphids and parasitoid wasps was reduced towards the middle of all fields while aphid symbiont species richness and diversity decreased into the field in conventional, but not integrated, field-strips. The proportion of aphids hosting different symbiont combinations varied across cropping systems, with distances into the fields, and were correlated with parasitoid wasp abundances. 4. In the pot experiment, aphids hosted up to six symbionts. Flower presence increased natural enemy abundance and diversity, and decreased aphid abundance. The proportion of aphids hosting different symbiont combinations varied across the flower treatment and time, and were correlated with varying abundances of the different specialist parasitoid wasp species recruited by different flowers. 5. Synthesis and applications. Floral plantings and flower identity can have community-wide impacts on the combinations of bacterial endosymbionts hosted by herbivorous insects. Our work highlights the potential of within-season selection for symbiont-mediated pest resistance to natural enemies with biological control impacts. This could be mitigated through increased recruitment of diverse natural enemies by incorporating functional diversity of floral resources into the environment.
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