Joachim Töpper

From facilitation to competition: temperature-driven shift in dominant plant interactions affects population dynamics in seminatural grasslands.

Biotic interactions are often ignored in assessments of climate change impacts. However, climate-related changes in species interactions, often mediated through increased dominance of certain species or functional groups, may have important implications for how species respond to climate warming and altered precipitation patterns. We examined how a dominant plant functional group affected the population dynamics of four co-occurring forb species by experimentally removing graminoids in seminatural grasslands. Specifically, we explored how the interaction between dominants and subordinates varied with climate by replicating the removal experiment across a climate grid consisting of 12 field sites spanning broad-scale temperature and precipitation gradients in southern Norway. Biotic interactions affected population growth rates of all study species, and the net outcome of interactions between dominants and subordinates switched from facilitation to competition with increasing temperature along the temperature gradient. The impacts of competitive interactions on subordinates in the warmer sites could primarily be attributed to reduced plant survival. Whereas the response to dominant removal varied with temperature, there was no overall effect of precipitation on the balance between competition and facilitation. Our findings suggest that global warming may increase the relative importance of competitive interactions in seminatural grasslands across a wide range of precipitation levels, thereby favouring highly competitive dominant species over subordinate species. As a result, seminatural grasslands may become increasingly dependent on disturbance (i.e. traditional management such as grazing and mowing) to maintain viable populations of subordinate species and thereby biodiversity under future climates. Our study highlights the importance of population-level studies replicated under different climatic conditions for understanding the underlying mechanisms of climate change impacts on plants.

Management-driven evolution in a domesticated ecosystem

Millennia of human land-use have resulted in the widespread occurrence of what have been coined ‘domesticated ecosystems’. The anthropogenic imprints on diversity, composition, structure and functioning of such systems are well documented. However, evolutionary consequences of human activities in these ecosystems are enigmatic. Calluna vulgaris (L.) is a keystone species of coastal heathlands in northwest Europe, an ancient semi-natural landscape of considerable conservation interest. Like many species from naturally fire-prone ecosystems, Calluna shows smoke-adapted germination, but it is unclear whether this trait arose prior to the development of these semi-natural landscapes or is an evolutionary response to the anthropogenic fire regime. We show that smoke-induced germination in Calluna is found in populations from traditionally burnt coastal heathlands but is lacking in naturally occurring populations from other habitats with infrequent natural fires. Our study thus demonstrates evolutionary imprints of human land-use in semi-natural ecosystems. Evolutionary consequences of historic anthropogenic impacts on wildlife have been understudied, but understanding these consequences is necessary for informed conservation and ecosystem management.

Biotic interaction effects on seedling recruitment along bioclimatic gradients: testing the stress-gradient hypothesis

Questions Is there a shift from positive to negative biotic interaction effects on seedling recruitment along two different stress gradients, temperature and precipitation (the stress-gradient hypothesis); do such interaction effects differ between species with different bioclimatic affinities? Location Boreal, sub-alpine and alpine grassland in southern Norway. Methods We tested the stress-gradient hypothesis by comparing seedling recruitment in bare-ground gaps where vegetation has been removed vs in extant grassland vegetation in 12 boreal, sub-alpine and alpine grassland sites along a precipitation gradient. This was tested in (1) a seed-sowing experiment and (2) in naturally occurring recruitment of alpine, generalist and boreal species. Results Emergence of the sown alpine species was higher in the cold alpine than in the warmer sub-alpine sites, with no effects of precipitation or vegetation removal. The sown generalists also decreased in emergence towards warmer sites, whereas there was no effect of temperature on the sown boreal species. Vegetation removal, interacting with precipitation, increased the emergence of the generalist and boreal species sown at intermediate precipitation levels. In contrast, interactions between temperature and vegetation removal regulated the emergence of all groups of naturally occurring seedlings. Alpine and generalist species emerged at the highest rate in alpine sites, whereas boreal species had highest emergence in the lowlands. Conclusion For all species groups, strong effects of vegetation removal show that competition from the extant vegetation dominates in controlling seedling emergence across all study sites and species. In generalist and boreal species, positive interactions between vegetation removal and temperature show that competitive interactions affect seedling emergence more strongly towards warmer climates, in line with the stress-gradient hypothesis. In contrast, alpine species show no such interactions. This suggests that species’ adaptations to climate, in combination with environmental forcing, control seedling emergence along the bioclimatic gradients. Our results have implications for nature conservation, as we propose that disturbance from grazing animals can be useful to release competition and thereby increase seedling recruitment and biodiversity in boreal and alpine grasslands in a warmer future.

The Response of Heterotrophic Prokaryote and Viral Communities to Labile Organic Carbon Inputs Is Controlled by the Predator Food Chain Structure

Factors controlling the community composition of marine heterotrophic prokaryotes include organic-C, mineral nutrients, predation, and viral lysis. Two mesocosm experiments, performed at an Arctic location and bottom-up manipulated with organic-C, had very different results in community composition for both prokaryotes and viruses. Previously, we showed how a simple mathematical model could reproduce food web level dynamics observed in these mesocosms, demonstrating strong top-down control through the predator chain from copepods via ciliates and heterotrophic nanoflagellates. Here, we use a steady-state analysis to connect ciliate biomass to bacterial carbon demand. This gives a coupling of top-down and bottom-up factors whereby low initial densities of ciliates are associated with mineral nutrient-limited heterotrophic prokaryotes that do not respond to external supply of labile organic-C. In contrast, high initial densities of ciliates give carbon-limited growth and high responsiveness to organic-C. The differences observed in ciliate abundance, and in prokaryote abundance and community composition in the two experiments were in accordance with these predictions. Responsiveness in the viral community followed a pattern similar to that of prokaryotes. Our study provides a unique link between the structure of the predator chain in the microbial food web and viral abundance and diversity.

How to induce defense responses in wild plant populations? Using bilberry (Vaccinium myrtillus) as example

Abstract Inducible plant defense is a beneficial strategy for plants, which imply that plants should allocate resources from growth and reproduction to defense when herbivores attack. Plant ecologist has often studied defense responses in wild populations by biomass clipping experiments, whereas laboratory and greenhouse experiments in addition apply chemical elicitors to induce defense responses. To investigate whether field ecologists could benefit from methods used in laboratory and greenhouse studies, we established a randomized block‐design in a pine‐bilberry forest in Western Norway. We tested whether we could activate defense responses in bilberry (Vaccinium myrtillus) by nine different treatments using clipping (leaf tissue or branch removal) with or without chemical treatment by methyljasmonate (MeJA). We subsequently measured consequences of induced defenses through vegetative growth and insect herbivory during one growing season. Our results showed that only MeJA‐treated plants showed consistent defense responses through suppressed vegetative growth and reduced herbivory by leaf‐chewing insects, suggesting an allocation of resources from growth to defense. Leaf tissue removal reduced insect herbivory equal to the effect of the MeJa treatments, but had no negative impact on growth. Branch removal did not reduce insect herbivory or vegetative growth. MeJa treatment and clipping combined did not give an additional defense response. In this study, we investigated how to induce defense responses in wild plant populations under natural field conditions. Our results show that using the chemical elicitor MeJA, with or without biomass clipping, may be a better method to induce defense response in field experiments than clipping of leaves or branches that often has been used in ecological field studies.

Advancing restoration ecology: A new approach to predict time to recovery

Handling Editor: Lars Brudvig Abstract 1. Species composition is a vital attribute of any ecosystem. Accordingly, ecological restoration often has the original, or “natural,” species composition as its target. However, we still lack adequate methods for predicting the expected time to compositional recovery in restoration studies. 2. We describe and explore a new, ordination regression-based approach (ORBA) for predicting time to recovery that allows both linear and asymptotic (logarithmic) relationships of compositional change with time. The approach uses distances between restored plots and reference plots along the successional gradient, represented by a vector in ordination space, to predict time to recovery. Thus, the approach rests on three requirements: (a) the general form of the relationship between compositional change and time must be known; (b) a sufficiently strong successional gradient must be present and adequately represented in a species compositional dataset; and (c) a restoration target must be specified. We tested the approach using data from a boreal old-growth forest that was followed for 18 years after experimental disturbance. Data from the first 9 years after disturbance were used to develop models, the subsequent 9 years for validation. 3. Rates of compositional recovery in the example dataset followed the general pattern of decrease with time since disturbance. Accordingly, linear models were too optimistic about the time to recovery, whereas the asymptotic models provided more precise predictions. 4. Synthesis and applications. Our results demonstrate that the new approach opens for reliable prediction of recovery rates and time to recovery using species compositional data. Moreover, it allows us to assess whether recovery proceeds in the desired direction and to quantitatively compare restoration speed, and hence effectiveness, between alternative management options.

Plastic Population Effects and Conservative Leaf Traits in a Reciprocal Transplant Experiment Simulating Climate Warming in the Himalayas

Climate warming poses considerable challenges for alpine plant species, especially for competitively inferior ones with resource-conservative adaptations to cold climates. The Himalayas are warming at rates considerably faster than the global average, so it is particularly important to assess how and through which mechanisms alpine plant species are affected there. We employed a demographic approach in a climate change experiment, where vegetation turfs were transplanted reciprocally between the central parts of the study species’ (Viola biflora L. var. rockiana) range and the warmer range margin, with a temperature difference of ca. 1°C. In addition, turfs were also transplanted outside the range to warmer habitats, simulating two different scenarios of climate warming, +1 and +4°C. Transplanting to warmer sites negatively impacted population growth rates (λ), survival and clonality, but did not affect growth and fecundity, while the productivity of the plant community increased. The reciprocal transplants to the colder habitat showed the opposite effects, for both V. biflora and the plant community, indicating plastic responses of the study species, driven by changes in plant–plant competition. However, the leaf traits underlying the modeled population growth rates were origin-site specific and not affected by the climate-change treatments over the study period, suggesting local adaptation of growth form to competition in the warmer range margin, and to climate adversity in the colder range center. The transplants outside the present species’ range showed consistently stronger reductions in population growth rate and survival, with mortality of 90–100% in the +4°C treatment. This illustrates that climatic changes beyond species’ present climatic ranges pose a serious risk for range contraction and extinction for Himalayan alpine species in the near future. As V. biflora seems mostly limited by competition under warming, its persistence in a future climate may become increasingly dependent on keeping competitive effects from the surrounding community low, for instance by management interventions like grazing and mowing.

Multiannual effects of induced plant defenses: Are defended plants good or bad neighbors?

Abstract Defenses induced by herbivore feeding or phytohormones such as methyl jasmonate (MeJA) can affect growth, reproduction, and herbivory, not only on the affected individual but also in its neighboring plants. Here, we report multiannual defense, growth, and reproductive responses of MeJA‐treated bilberry (Vaccinium myrtillus) and neighboring ramets. In a boreal forest in western Norway, we treated bilberry ramets with MeJA and water (control) and measured responses over three consecutive years. We observed the treatment effects on variables associated with herbivory, growth, and reproduction in the MeJA‐treated and untreated ramet and neighboring ramets distanced from 10 to 500 cm. MeJA‐treated ramets had fewer grazed leaves and browsed shoots compared to control, with higher effects in 2014 and 2015, respectively. In 2013, growth of control ramets was greater than MeJA‐treated ramets. However, MeJA‐treated ramets had more flowers and berries than control ramets 2 years after the treatment. The level of insect and mammalian herbivory was also lower in untreated neighboring ramets distanced 10–150 cm and, consistent with responses of MeJA‐treated ramets, the stronger effect was also one and 2 years delayed, respectively. The same neighboring ramets had fewer flowers and berries than untreated ramets, indicating a trade‐off between defense and reproduction. Although plant–plant effects were observed across all years, the strength varied by the distance between the MeJA‐treated ramets and its untreated neighbors. We document that induced defense in bilberry reduces both insect and mammalian herbivory, as well as growth, over multiple seasons. The defense responses occurred in a delayed manner with strongest effects one and 2 years after the induction. Additionally, our results indicate defense signaling between MeJA‐treated ramets and untreated neighbors. In summary, this study shows that induced defenses are important ecological strategies not only for the induced individual plant but also for neighboring plants across multiple years in boreal forests.