211. Root-feeding insect herbivores are of substantial evolutionary, ecological and economical 22 importance. Plants can resist insect herbivores through a variety of tolerance and resistance 23 strategies. To date, few studies have systematically assessed the prevalence and importance 24 of these strategies for root-herbivore interactions across different plant species. 25 2. Here, we characterize the defense strategies used by three different grassland species to 26 cope with a generalist root herbivore, the larvae of the European cockchafer Melolontha 27 melolontha. 28 3. Our results reveal that the different plant species rely on distinct sets of defense strategies.
29The spotted knapweed (Centaurea stoebe) resists attack by dissuading the larvae through 30 the release of repellent chemicals. White clover (Trifolium repens) does not repel the 31 herbivore, but reduces feeding, most likely through structural defenses and low nutritional 32 quality. Finally, the common dandelion (Taraxacum officinale) allows M. melolontha to 33 feed abundantly but compensates for tissue loss through induced regrowth. 34 4. Synthesis: Three co-occurring plant species have evolved different solutions to defend 35 themselves against attack by a generalist root herbivore. The different root defense 36 strategies may reflect distinct defense syndromes. 37 38 Keywords: belowground herbivores, chemical and structural defenses, generalist herbivores, 39 host resistance and tolerance, plant -insect interactions 40 65 4 (Agrawal & Fishbein, 2006) is an important next step towards a better understanding of the 66 ecology and evolution of root-herbivore interactions. 67In the present study, we combine different experimental approaches to understand the root-68 defense strategies of three different, co-occurring European grassland species: the common 69 dandelion Taraxacum officinale agg. (Asteraceae), the spotted knapweed Centaurea stoebe 70 (Asteraceae) and white clover Trifolium repens (Fabaceae). All three species co-occur with a 71 generalist root herbivore, the larva of the European cockchafer Melolontha melolontha 72 (Coleoptera: Scarabeidae). Melolontha melolontha is native to Europe and occurs abundantly 73 in grasslands. Its larvae develop best on this species (Hauss, 1975;Hauss & Schütte, 1976). The 74 reasons for this preference and host suitability are unknown. Recently, it was shown that C. 75 stoebe is a bad host for M. melolontha larvae (Huang, Zwimpfer, Hervé, Bont, & Erb, 2018). 76 The host suitability of T. repens is less clear (Huang et al., 2018; Sukovata, Jaworski, 77 Karolewski, & Kolk, 2015). Regarding potential defense strategies of the three species against 78 root-herbivores, mechanistic work so far has mostly focused on T. officinale. Upon damage, T. 79 officinale releases a bitter latex sap containing high amount of the sesquiterpene lactone 80 taraxinic acid β-D-glucopyranosyl ester (TA-G) (Huber et al., 2015). High TA-G levels are 81 associated with reduced M. melolontha damage, and silencing TA-G produc...