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Current knowledge about defense strategies in plants under herbivore pressure is predominantly based on vascular plants. Bryophytes are rarely consumed by herbivores since they have ample secondary metabolites. However, it is unknown whether bryophytes have induced defenses against herbivory and whether there is a trade-off between growth and defense in bryophytes. METHODS:In an experiment with two peatland bryophytes, Sphagnum magellanicum Brid. and S. fuscum (Schimp.) H. Klinggr., two kinds of herbivory, clipping with scissors and grazing by mealworms (Tenebrio molitor L.) were simulated. At the end of the experiment, we measured growth traits, carbon-based defense compounds (total phenolics and cellulose) and storage compounds (total nonstructural carbohydrates) of these two Sphagnum species.RESULTS: Grazing but not clipping increased total phenolics and C:N ratio and reduced biomass production and height increment. A negative relationship between biomass production and total phenolics was found in S. magellanicum but not in S. fuscum, indicating a growth-defense trade-off that is species-specific. Grazing reduced the sugar starch content of S. magellanicum and the sugar of S. fuscum. Either clipping or grazing had no effect on chlorophyll fluorescence (including actual and maximum photochemical efficiency of photosystem II) except that a significant effect of clipping on actual photochemical efficiency in S. fuscum was observed. CONCLUSIONS:Our results suggest that Sphagnum can have induced defense against herbivory and that this defense can come at a cost of growth. These findings advance our knowledge about induced defense in bryophytes, the earliest land plants.

In a natural environment, plants usually interact with their neighbors predominantly through resource competition, allelopathy, and facilitation. The occurrence of the positive effect of allelopathy between peat mosses (Sphagnum L.) is rare, but it has been observed in a field experiment. It is unclear whether the stability of the water table level in peat induces positive vs. negative effects of allelopathy and how that is related to phenolic allelochemical production in Sphagnum. Based on field experiment data, we established a laboratory experiment with three neighborhood treatments to measure inter-specific interactions between Sphagnum angustifolium (Russ.) C. Jens and Sphagnum magellanicum Brid. We found that the two species were strongly suppressed by the allelopathic effects of each other. S. magellanicum allelopathically facilitated S. angustifolium in the field but inhibited it in the laboratory, and relative allelopathy intensity appeared to be positively related to the content of released phenolics. We conclude that the interaction type and intensity between plants are dependent on environmental conditions. The concentration of phenolics alone may not explain the type and relative intensity of allelopathy. Carefully designed combined field and laboratory experiments are necessary to reveal the mechanism of species interactions in natural communities.