Abstract. Phenological mismatches due to climate change may have important ecological consequences. In a three-year study, phenological shifts due to experimental warming markedly altered trophic relationships between plants and insect herbivores, causing a dramatic decline of reproductive capacity for one of the plant species. In a Tibetan meadow, the gentian (Gentiana formosa) typically flowers after the peak larva density of a noctuid moth (Melanchra pisi ) that primarily feeds on a dominant forb (anemone, Anemone trullifolia var. linearis). However, artificial warming of ;1.58C advanced gentian flower phenology and anemone vegetative phenology by a week, but delayed moth larvae emergence by two weeks. The warming increased larval density 10-fold, but decreased anemone density by 30%. The phenological and density shifts under warmed conditions resulted in the insect larvae feeding substantially on the gentian flowers and ovules; there was ;100-fold more damage in warmed than in unwarmed chambers. This radically increased trophic connection reduced gentian plant reproduction and likely contributed to its reduced abundance in the warmed chambers.

Summary• Temperature is projected to increase more during the winter than during the summer in cold regions. The effects of winter warming on reproductive effort have not been examined for temperate plant species.• Here, we report the results of experimentally induced seasonal winter warming (0.4 and 2.4°C increases in growing and nongrowing seasons, respectively, using warmed and ambient open-top chambers in a Tibetan Plateau alpine meadow) for nine indeterminate-growing species producing multiple (single-flowered or multi-flowered) inflorescences and three determinate-growing species producing single inflorescences after a 3-yr period of warming.• Warming reduced significantly flower number and seed production per plant for all nine multi-inflorescence species, but not for the three single-inflorescence species. Warming had an insignificant effect on the fruit to flower number ratio, seed size and seed number per fruit among species. The reduction in seed production was largely attributable to the decline in flower number per plant. The flowering onset time was unaffected for nine of the 12 species. Therefore, the decline in flower production and seed production in response to winter warming probably reflects a physiological response (e.g. metabolic changes associated with flower production).• Collectively, the data indicate that global warming may reduce flower and seed production for temperate herbaceous species and will probably have a differential effect on single-vs multi-inflorescence species.

Summary 1.It is well known that climate change alters abiotic factors (temperature and water availability) that directly affect ecosystem properties. However, less is known about the indirect impacts of climate change on ecosystem structure and function. Here, we show that experimental warming may deteriorate ecosystems via trophic interactions. 2. In a Tibetan alpine meadow, plant species composition, size, coverage and above-ground biomass were investigated to reveal the effect of artificial warming (c. 1°C mean annual temperature at the soil surface), which was accomplished using warmed and ambient open top chambers. In addition, rodent damage to plants was assessed. 3. The dicot forb silverweed Potentilla anserina increased significantly, while other species groups remained unchanged or decreased in plant community dominance rank after 2 years of artificial warming. The change in community structure was attributed to the difference in biomass allocation and growth form among species. 4. In the third year, plateau zokors Myospalax fontanierii, a widespread rodent herbivore, damaged plants in the warmed chambers, while leaving plants in the ambient chambers mostly undamaged. Above-ground biomass was found to be smaller in the warmed chambers than the controls in the third year, in contrast to the trend of the first 2 years. In addition, zokor burrow density was positively correlated with silverweed biomass and its dominance within communities, which was consistent with findings of independent field investigations that silverweed-dominated plots were more likely to be visited and damaged by the zokors than sites-dominated by grass species. 5. Synthesis and applications. The top-down negative effect of zokor damage on above-ground biomass in the warmed chambers was induced by the bottom-up effect of changes in species composition and community structure on zokor foraging behaviour, which were driven by artificial warming. Such trophic interactions may invalidate some predictions of ecological effects by current species-climate envelope models. Furthermore, because management measures including increasing the water table, planting grass and moderate cattle grazing may prevent silverweed dominance, we suggest that these interventions could be employed to control zokor damage in alpine meadows that are predicted to be drier and warmer in the future.