Ciska Veen

Dr. Ciska Veen



Droevendaalsesteeg 10
6708 PB Wageningen

+31 (0) 317 47 34 00

The Netherlands



Soil biodiversity is at the basis of all life on earth. Understanding the functions performed by soil organisms is essential to sustainable use of soils.


Ciska Veen obtained her MSc (2005) and PhD (2011) degree at the University of Groningen (the Netherlands), where she has been studying how grazing by large vertebrate herbivores affects soil biodiversity and ecosystem functioning in grasslands. She then obtained an Rubicon (2011; SLU, Sweden) and Veni (2014; NIOO-KNAW) grant to untangle how microbial communities drive soil carbon and nutrient cycling. In 2020, Ciska started as a tenure-track researcher (funded by an Aspasia grant) at the department of Terrestrial Ecology at the Netherlands Institute of Ecology. In her current research she aims at understanding how soil biodiversity drives soil functioning and carbon storage and how we can steer soil communities for sustainable land-use and climate change mitigation. She is currently leading a research project on how biodiversity and ecosystem functioning is changing when agricultural land is transformed into food forests and on the role of soils in climate-smart forest management. During her career Ciska has actively contributed to building an inclusive scientific community.




  • 2020–Present
    Tenure track researcher at (NIOO-KNAW)
  • 2018–2020
    Junior Group leader (NIOO-KNAW)
  • 2014–2018
    Postdoctoral researcher (NIOO-KNAW, NWO-VENI)
  • 2011–2014
    Postdoctoral researcher (SLU, Sweden, NWO-Rubicon)
  • 2011
    Postdoctoral researcher Aquatic Ecology (NIOO-KNAW)


  • 2005–2011
    PhD student (University of Groningen)
  • 1999–2005
    BSc and MSc (University of Groningen)

Editorial board memberships

  • 2019–Present
    Editor at Functional Ecology
  • 2020–Present
    Editor at Oikos

PhD students

  • 2021–Present
    Steven de Goede
    Promotors en Copromotors: Ciska Veen, Emilia Hannula, Frank Sterck, Wim van der Putten
  • 2020–Present
    Isabelle van der Zanden
    Promotors en Copromotors: Ciska Veen, Wim van der Putten
  • 2020–Present
    Zhaoqi Bin
    Promotors en Copromotors: Wim van der Putten, Ciska Veen
  • 2020–Present
    Felipe Cozim Melges
    Wageningen University; NIOO-KNAW
    Promotors en Copromotors: Hannah van Zanten, Raimon Ripoll Bosch, Ciska Veen
  • 2018–Present
    Keli Li
    Promotors en Copromotors: Wim van der Putten, Ciska Veen
  • 2017–Present
    Sophie van Rijssel
    Promotors en Copromotors: Wim van der Putten, Ciska Veen
  • 2014–2018
    Marta Manrubia
    Promotors en Copromotors: Wim van der Putten, Ciska Veen

Invited talks and keynote addresses on symposia and conferences

  • 2022
    Above-belowground interactions BES-PSI
  • 2021
    HSTalk Plant-soil feedback
  • 2019
    Plant-soil feedback symposium


Peer-reviewed publicaties

  • Biogeochemistry

    Coarse woody debris accelerates the decomposition of deadwood inputs across temperate forest

    Mark Bradford, Ciska Veen, Ella M. Bradford, Kristofer R. Covey, Tom Crowther, Nicholas Fields, Paul T. Frankson, Javier González-Rivero, Fiona V. Jevon, Sara E. Kuebbing, Steven McBride, Jacqueline E. Mohan, Emily E. Oldfield, Angela M. Oliverio, Alexander Polussa, Corinna Steinrueck, Michael S. Strickland, Elisabeth B. Ward, Carl Wepking, Daniel S. Maynard
    Wood decomposition is regulated by multiple controls, including climate and wood traits, that vary at local to regional scales. Yet decomposition rates differ dramatically when these controls do not. Fungal community dynamics are often invoked to explain these differences, suggesting that knowledge of ecosystem properties that influence fungal communities will improve understanding and projection of wood decomposition. We hypothesize that deadwood inputs decompose faster in forests with higher stocks of downed coarse woody material (CWM) because CWM is a resource from which lignocellulolytic fungi rapidly colonize new inputs. To test this hypothesis, we measure decomposition of 1,116 pieces of fine woody material (FWM) of five species, incubated for 13 to 49 months at five locations spanning 10°-latitude in eastern U.S. forest. We place FWM pieces near and far from CWM across observational transects and experimental common gardens. Soil temperature positively affects location-level mean decomposition rates, but these among-location differences are smaller than within-location variation in decomposition. Some of this variability is caused by CWM, where FWM pieces next to CWM decompose more rapidly. These effects are greater with time of incubation and lower initial wood density of FWM. The effect size of CWM is of the same relative magnitude as for the known controls of temperature, deadwood density and diameter. Abundance data for CWM is available for many forests and hence may be an ecosystem variable amenable for inclusion in decomposition models. Our findings suggest that conservation efforts to rebuild depleted CWM stocks in temperate forests may accelerate decomposition of fresh deadwood inputs.
  • Biotropica

    Soil nutrient dissimilarity and litter nutrient limitation as major drivers of home field advantage in riparian tropical forests

    Rebeca Leme Oliva, Ciska Veen, Marcel Okamoto Tanaka
    Decomposition is a key process driving carbon and nutrient cycling in ecosystems worldwide. The home field advantage effect (HFA) has been found to accelerate decomposition rates when litter originates from “home” when compared to other (“away”) sites. It is still poorly known how HFA plays out in tropical, riparian forests, particularly in forests under restoration. We carried out three independent reciprocal litter transplant experiments to test how litter quality, soil nutrient concentrations, and successional stage (age) influenced HFA in tropical riparian forests. These experimental areas formed a wide gradient of soil and litter nutrients, which we used to evaluate the more general hypothesis that HFA varies with dissimilarity in soil nutrients and litter quality. We found that HFA increased with soil nutrient dissimilarity, suggesting that litter translocation uncouples relationships between decomposers and litter characteristics; and with litter N:P, indicating P limitation in this system. We also found negative HFA effects at a site under restoration that presented low decomposer ability, suggesting that forest restoration does not necessarily recover decomposer communities and nutrient cycling. Within each of the independent experiments, the occurrence of HFA effects was limited and their magnitude was not related to forest age, nor soil and litter quality. Our results imply that HFA effects in tropical ecosystems are influenced by litter nutrient limitation and soil nutrient dissimilarity between home and away sites, but to further disentangle major HFA drivers in tropical areas, a gradient of dissimilarity between litter and soil properties must be implemented in future experimental designs.
  • Nature Communications

    Environmental heterogeneity modulates the effect of plant diversity on the spatial variability of grassland biomass

    Pedro Daleo, Juan Alberti, Enrique J. Chaneton, Oscar Iribarne, Pedro M. Tognetti, Jonathan D. Bakker, Elizabeth T. Borer, Martín Bruschetti, Andrew S. MacDougall, Jesús Pascual, Mahesh Sankaran, Eric W. Seabloom, Shaopeng Wang, Sumanta Bagchi, Lars A. Brudvig, Jane A. Catford, Chris R. Dickman, Timothy L. Dickson, Ian Donohue, Nico Eisenhauer, Daniel S. Gruner, Sylvia Haider, Anke Jentsch, Johannes M. H. Knops, Ylva Lekberg, Rebecca L. McCulley, Joslin L. Moore, Brent Mortensen, Timothy Ohlert, Meelis Pärtel, Pablo L. Peri, Sally A. Power, Anita C. Risch, Camila Rocca, Nicholas G. Smith, Carly J. Stevens, Riin Tamme, Ciska Veen, Peter A. Wilfahrt, Yann Hautier

    Plant productivity varies due to environmental heterogeneity, and theory suggests that plant diversity can reduce this variation. While there is strong evidence of diversity effects on temporal variability of productivity, whether this mechanism extends to variability across space remains elusive. Here we determine the relationship between plant diversity and spatial variability of productivity in 83 grasslands, and quantify the effect of experimentally increased spatial heterogeneity in environmental conditions on this relationship. We found that communities with higher plant species richness (alpha and gamma diversity) have lower spatial variability of productivity as reduced abundance of some species can be compensated for by increased abundance of other species. In contrast, high species dissimilarity among local communities (beta diversity) is positively associated with spatial variability of productivity, suggesting that changes in species composition can scale up to affect productivity. Experimentally increased spatial environmental heterogeneity weakens the effect of plant alpha and gamma diversity, and reveals that beta diversity can simultaneously decrease and increase spatial variability of productivity. Our findings unveil the generality of the diversity-stability theory across space, and suggest that reduced local diversity and biotic homogenization can affect the spatial reliability of key ecosystem functions.
  • Microbiome

    Mycorrhiza-mediated recruitment of complete denitrifying Pseudomonas reduces N2O emissions from soil

    Xia Li, Ruotong Zhao, Dandan Li, Guangzhou Wang, Shuikuan Bei, Xiaotang Ju, Ran Nathan, Long Li, Thomas W. Kuyper, Peter Christie, Franz S. Bender, Ciska Veen, Marcel G. A. van der Heijden, Wim H. van der Putten, Fusuo Zhang, Klaus Butterbach-Bahl, Junling Zhang
    Arbuscular mycorrhizal fungi (AMF) are key soil organisms and their extensive hyphae create a unique hyphosphere associated with microbes actively involved in N cycling. However, the underlying mechanisms how AMF and hyphae‑associated microbes may cooperate to influence N2O emissions from “hot spot” residue patches remain unclear. Here we explored the key microbes in the hyphosphere involved in N 2O production and consumption using amplicon and shotgun metagenomic sequencing. Chemotaxis, growth and N2O emissions of isolated N2O‑reducing bacteria in response to hyphal exudates were tested using in vitro cultures and inoculation experiments.

    AMF hyphae reduced denitrification‑derived N 2O emission (max. 63%) in C‑ and N‑rich residue patches. AMF consistently enhanced the abundance and expression of clade I nosZ gene, and inconsistently increased that of nirS and nirK genes. The reduction of N2 O emissions in the hyphosphere was linked to N2O‑reducing Pseudomonas specifically enriched by AMF, concurring with the increase in the relative abundance of the key genes involved in bacterial citrate cycle. Phenotypic characterization of the isolated complete denitrifying P. fluorescens strain JL1 (possessing clade I nosZ) indicated that the decline of net N 2 O emission was a result of upregulated nosZ expression in P. fluorescens following hyphal exudation (e.g. carboxylates). These findings were further validated by re‑inoculating sterilized residue patches with P. fluorescens and by an 11‑year‑long field experiment showing significant positive correlation between hyphal length density with the abundance of clade I nosZ gene.

    The cooperation between AMF and the N2O‑reducing Pseudomonas residing on hyphae significantly reduce N2O emissions in the microsites. Carboxylates exuded by hyphae act as attractants in recruiting P. fluorescens and also as stimulants triggering nosZ gene expression. Our discovery indicates that reinforcing synergies between AMF and hyphosphere microbiome may provide unexplored opportunities to stimulate N 2 O consumption in nutrient‑enriched microsites, and consequently reduce N2O emissions from soils. This knowledge opens novel avenues to exploit cross‑kingdom microbial interactions for sustainable agriculture and for climate change mitigation.
  • Ecological Entomology

    Contrasting effects of nitrogen fertiliser application on the performance of closely related grasshoppers through changes in plant nutrient concentrations

    Yu Zhu, Quanhui Ma, Zhiwei Zhong, Ming Jiang, (Liesbeth) E.S. Bakker, Jeff A. Harvey, Ciska Veen, Cong Chen, Deli Wang
    Global environmental changes mediated by anthropogenic processes can affect the nutrient status of plants, with important consequences for the performance and dynamics of insect herbivores that feed on them.
    While it is well documented that insects from different feeding guilds (e.g., sap-feeders and leaf-chewers) can respond differently to altered food resources due to their distinct physiological and ecological characteristics, little is known about how ecologically similar insect species from the same feeding guild respond to changes in food nutrient status.
    Using nitrogen (N) fertiliser, the authors examined the effects of N inputs on two sympatric grasshopper species, Euchorthippus cheui and E. unicolor, that share the same host food plant, Leymus chinensis grass. The authors examined the effects of fertilisation on the individual feeding behaviour, performance and abundance of the two grasshopper species.
    The nutrient (protein) content of L. chinensis leaves was enhanced by fertilisation during the entire season. However, E. cheui and E. unicolor exhibited differing growth rates, development and body size responses to fertilisation.
    E. cheui preferred L. chinensis leaves from high-N fertilised treatments, while E. unicolor preferred leaves from low-N fertilised treatments. Moreover, fertilisation increased the abundance of E. cheui but had no significant effect on the abundance of E. unicolor in the field.
    The findings imply that effective management and conservation strategies for insects should target the needs of individual species rather than species groups or communities as a whole because nutritional and environmental requirements are often species-specific.
  • Ecology Letters

    Soil legacy effects of plants and drought on aboveground insects in native and range-expanding plant communities

    Soils contain biotic and abiotic legacies of previous conditions that may influence plant community biomass and associated aboveground biodiversity. However, little is known about the relative strengths and interactions of the various belowground legacies on aboveground plant–insect interactions. We used an outdoor mesocosm experiment to investigate the belowground legacy effects of range-expanding versus native plants, extreme drought and their interactions on plants, aphids and pollinators. We show that plant biomass was influenced more strongly by the previous plant community than by the previous summer drought. Plant communities consisted of four congeneric pairs of natives and range expanders, and their responses were not unanimous. Legacy effects affected the abundance of aphids more strongly than pollinators. We conclude that legacies can be contained as soil ‘memories’ that influence aboveground plant community interactions in the next growing season. These soil-borne ‘memories’ can be altered by climate warming-induced plant range shifts and extreme drought.
  • Ecology Letters

    Linking changes in species composition and biomass in a globally distributed grassland experiment

    Emma Ladouceur, Shane A. Blowes, Jonathan M. Chase, Adam T. Clark, Magda Garbowski, Juan Alberti, Carlos Alberto Arnillas, Jonathan D. Bakker, Isabel C. Barrio, Siddharth Bharath, Elizabeth T. Borer, Lars A. Brudvig, Marc W. Cadotte, Qingqing Chen, Scott L. Collins, Chris R. Dickman, Ian Donohue, Guozhen Du, Anne Ebeling, Nico Eisenhauer, Philip A. Fay, Nicole Hagenah, Yann Hautier, Anke Jentsch, I.S. Jonsdottir, Kimberly Komatsu, Andrew S. MacDougall, Jason P. Martina, Joslin L. Moore, John W Morgan, Pablo L. Peri, Sally A. Power, Zhengwei Ren, Anita C. Risch, Christiane Roscher, Max A. Schuchardt, Eric W. Seabloom, Carly J. Stevens, Ciska Veen, Risto Virtanen, Glenda M. Wardle, Peter A. Wilfahrt, W. Stanley Harpole

    Global change drivers, such as anthropogenic nutrient inputs, are increasing globally. Nutrient deposition simultaneously alters plant biodiversity, species composition and ecosystem processes like aboveground biomass production. These changes are underpinned by species extinction, colonisation and shifting relative abundance. Here, we use the Price equation to quantify and link the contributions of species that are lost, gained or that persist to change in aboveground biomass in 59 experimental grassland sites. Under ambient (control) conditions, compositional and biomass turnover was high, and losses (i.e. local extinctions) were balanced by gains (i.e. colonisation). Under fertilisation, the decline in species richness resulted from increased species loss and decreases in species gained. Biomass increase under fertilisation resulted mostly from species that persist and to a lesser extent from species gained. Drivers of ecological change can interact relatively independently with diversity, composition and ecosystem processes and functions such as aboveground biomass due to the individual contributions of species lost, gained or persisting.
  • Ecological Monographs

    Temporal dynamics of range-expander and congeneric native plant responses during and after extreme drought events

    Qiang Yang, Ciska Veen, Roel Wagenaar, Marta Manrubia-Freixa, Freddy ten Hooven, Wim H. van der Putten

    Climate change is causing range shifts of many species to higher latitudes and altitudes and increasing their exposure to extreme weather events. It has been shown that range-shifting plant species may perform differently in new soil than related natives; however, little is known about how extreme weather events affect range-expanding plants compared to related natives. In this study we used outdoor mesocosms to study how range-expanding plant species responded to extreme drought in live soil from a habitat in a new range with and without live soil from a habitat in the original range (Hungary). During summer drought, the shoot biomass of the range-expanding plant community declined. In spite of this, in the mixed community, range expanders produced more shoot biomass than congeneric natives. In mesocosms with a history of range expanders in the previous year, native plants produced less biomass. Plant legacy or soil origin effects did not change the response of natives or range expanders to summer drought. During rewetting, range expanders had less biomass than congeneric natives but higher drought resilience (survival) in soils from the new range where in the previous year native plant species had grown. The biomass patterns of the mixed plant communities were dominated by Centaurea spp.; however, not all plant species within the groups of natives and of range expanders showed the general pattern. Drought reduced the litter decomposition, microbial biomass, and abundances of bacterivorous, fungivorous, and carnivorous nematodes. Their abundances recovered during rewetting. There was less microbial and fungal biomass, and there were fewer fungivorous nematodes in soils from the original range where range expanders had grown in the previous year. We concluded that in mixed plant communities of range expanders and congeneric natives, range expanders performed better, under both ambient and drought conditions, than congeneric natives. However, when considering the responses of individual species, we observed variations among pairs of congenerics, so that under the present mixed-community conditions there was no uniformity in responses to drought of range expanders versus congeneric natives. Range-expanding plant species reduced soil fungal biomass and the numbers of soil fungivorous nematodes, suggesting that the effects of range-expanding plant species can trickle up in the soil food web.
  • Science

    Termite sensitivity to temperature affects global wood decay rates

    Amy E. Zanne, Habacuc Flores-Moreno, Jeff R. Powell, William K. Cornwell, James W. Dalling, Amy T. Austin, Aimée T. Classen, Paul Eggleton, Kei Ichi Okada, Catherine L. Parr, E. Carol Adair, Stephen Adu-Bredu, Md Azharul Alam, Carolina Alvarez-Garzón, Deborah Mattos Guimarães Apgaua, Roxana Aragón, Marcelo Ardon, Stefan K. Arndt, Louise A. Ashton, Nicholas A. Barber, Jacques Beauchêne, Matty P. Berg, Jason Beringer, Matthias M. Boer, José Antonio Bonet, Katherine Bunney, Tynan J. Burkhardt, Dulcinéia Carvalho, Dennis Castillo-Figueroa, Lucas A. Cernusak, Alexander W. Cheesman, Tainá M. Cirne-Silva, Jamie R. Cleverly, Johannes H. C. Cornelissen, Timothy J. Curran, André M. D’Angioli, Caroline Dallstream, Nico Eisenhauer, Fidele Evouna Ondo, Alex Fajardo, Romina D. Fernandez, Astrid Ferrer, Marco A.L. Fontes, Mark L. Galatowitsch, Grizelle González, Felix Gottschall, Peter R. Grace, Elena Granda, Hannah M. Griffiths, Mariana Guerra Lara, Motohiro Hasegawa, Mariet M. Hefting, Nina Hinko-Najera, Lindsay B. Hutley, Jennifer Jones, Anja Kahl, Mirko Karan, Joost Keuskamp, Tim Lardner, Michael Liddell, Craig Macfarlane, Cate Macinnis-Ng, Ravi F. Mariano, M. Soledad Méndez, Wayne S. Meyer, Akira S. Mori, Aloysio S. Moura, Matthew Northwood, Romà Ogaya, Rafael S. Oliveira, Alberto Orgiazzi, Juliana Pardo, Guille Peguero, Josep Peñuelas, Luis I. Perez, Juan M. Posada, Cecilia M. Prada, Tomáš Přívětivý, Suzanne M. Prober, Jonathan Prunier, Gabriel W. Quansah, Víctor Resco de Dios, Ronny Richter, Mark P. Robertson, Lucas F. Rocha, Megan A. Rúa, Carolina Sarmiento, Richard P. Silberstein, Mateus C. Silva, Flávia Freire Siqueira, Matthew Glenn Stillwagon, Jacqui Stol, Melanie K. Taylor, François P. Teste, David Y.P. Tng, David Tucker, Manfred Türke, Michael D. Ulyshen, Oscar J. Valverde-Barrantes, Eduardo van den Berg, Richard S.P. van Logtestijn, Ciska Veen, Jason G. Vogel, Timothy J. Wardlaw, Georg Wiehl, Christian Wirth, Michaela J. Woods, Paul Camilo Zalamea

    Deadwood is a large global carbon store with its store size partially determined by biotic decay. Microbial wood decay rates are known to respond to changing temperature and precipitation. Termites are also important decomposers in the tropics but are less well studied. An understanding of their climate sensitivities is needed to estimate climate change effects on wood carbon pools. Using data from 133 sites spanning six continents, we found that termite wood discovery and consumption were highly sensitive to temperature (with decay increasing >6.8 times per 10°C increase in temperature)—even more so than microbes. Termite decay effects were greatest in tropical seasonal forests, tropical savannas, and subtropical deserts. With tropicalization (i.e., warming shifts to tropical climates), termite wood decay will likely increase as termites access more of Earth’s surface.
  • Molecular Ecology

    Soil microbial diversity and community composition during conversion from conventional to organic agriculture

    Sophie van Rijssel, Ciska Veen, Guusje Koorneef, Tanja Bakx-Schotman, Freddy ten Hooven, Stefan Geisen, Wim H. van der Putten
    It is generally assumed that the dependence of conventional agriculture on artificial fertilizers and pesticides strongly impacts the environment, while organic agriculture relying more on microbial functioning may mitigate these impacts. However, it is not well known how microbial diversity and community composition change in conventionally managed farmers' fields that are converted to organic management. Here, we sequenced bacterial and fungal communities of 34 organic fields on sand and marine clay soils in a time series (chronosequence) covering 25 years of conversion. Nearby conventional fields were used as references. We found that community composition of bacteria and fungi differed between organic and conventionally managed fields. In the organic fields, fungal diversity increased with time since conversion. However, this effect disappeared when the conventional paired fields were included. There was a relationship between pH and soil organic matter content and the diversity and community composition of bacteria and fungi. In marine clay soils, when time since organic management increased, fungal communities in organic fields became more dissimilar to those in conventional fields. We conclude that conversion to organic management in these Dutch farmers' fields did not increase microbial community diversity. Instead, we observed that in organic fields in marine clay when time since conversion increased soil fungal community composition became progressively dissimilar from that in conventional fields. Our results also showed that the paired sampling approach of organic and conventional fields was essential in order to control for environmental variation that was otherwise unaccounted for.
  • ISME Journal

    Microbial storage and its implications for soil ecology

    Kyle Mason-Jones, Serina L. Robinson, Ciska Veen, S. Manzoni, Wim H. van der Putten
    Organisms throughout the tree of life accumulate chemical resources, in particular forms or compartments, to secure their availability for future use. Here we review microbial storage and its ecological significance by assembling several rich but disconnected lines of research in microbiology, biogeochemistry, and the ecology of macroscopic organisms. Evidence is drawn from various systems, but we pay particular attention to soils, where microorganisms play crucial roles in global element cycles. An assembly of genus-level data demonstrates the likely prevalence of storage traits in soil. We provide a theoretical basis for microbial storage ecology by distinguishing a spectrum of storage strategies ranging from surplus storage (storage of abundant resources that are not immediately required) to reserve storage (storage of limited resources at the cost of other metabolic functions). This distinction highlights that microorganisms can invest in storage at times of surplus and under conditions of scarcity. We then align storage with trait-based microbial life-history strategies, leading to the hypothesis that ruderal species, which are adapted to disturbance, rely less on storage than microorganisms adapted to stress or high competition. We explore the implications of storage for soil biogeochemistry, microbial biomass, and element transformations and present a process-based model of intracellular carbon storage. Our model indicates that storage can mitigate against stoichiometric imbalances, thereby enhancing biomass growth and resource-use efficiency in the face of unbalanced resources. Given the central roles of microbes in biogeochemical cycles, we propose that microbial storage may be influential on macroscopic scales, from carbon cycling to ecosystem stability.
  • Nature Ecology and Evolution

    Evolutionary history of grazing and resources determine herbivore exclusion effects on plant diversity

    Jodi N. Price, Judith Sitters, Timothy Ohlert, Pedro M. Tognetti, Cynthia S. Brown, Eric W. Seabloom, Elizabeth T. Borer, Suzanne M. Prober, (Liesbeth) E.S. Bakker, Andrew S. MacDougall, Laura Yahdjian, Daniel S. Gruner, Harry Olde Venterink, Isabel C. Barrio, Pamela Graff, Sumanta Bagchi, Carlos Alberto Arnillas, Jonathan D. Bakker, Dana M. Blumenthal, Elizabeth H Boughton, Lars A. Brudvig, Miguel N. Bugalho, Marc W. Cadotte, Maria C. Caldeira, Chris R. Dickman, Ian Donohue, Sonnier Grégory, Yann Hautier, I.S. Jonsdottir, Luciola S. Lannes, Rebecca L. McCulley, Joslin L. Moore, Sally A. Power, Anita C. Risch, Martin Schütz, Rachel Standish, Carly J. Stevens, Ciska Veen, Risto Virtanen, Glenda M. Wardle

    Ecological models predict that the effects of mammalian herbivore exclusion on plant diversity depend on resource availability and plant exposure to ungulate grazing over evolutionary time. Using an experiment replicated in 57 grasslands on six continents, with contrasting evolutionary history of grazing, we tested how resources (mean annual precipitation and soil nutrients) determine herbivore exclusion effects on plant diversity, richness and evenness. Here we show that at sites with a long history of ungulate grazing, herbivore exclusion reduced plant diversity by reducing both richness and evenness and the responses of richness and diversity to herbivore exclusion decreased with mean annual precipitation. At sites with a short history of grazing, the effects of herbivore exclusion were not related to precipitation but differed for native and exotic plant richness. Thus, plant species’ evolutionary history of grazing continues to shape the response of the world’s grasslands to changing mammalian herbivory.
  • Global Change Biology

    Global maps of soil temperature

    Jonas J Lembrechts, Johan van den Hoogen, Juha Aalto, Michael B Ashcroft, Pieter De Frenne, Julia Kemppinen, Martin Kopecký, Miska Luoto, Ilya M. D. Maclean, Tom Crowther, Joseph J Bailey, Stef Haesen, David H Klinges, Pekka Niittynen, Brett R Scheffers, Koenraad Van Meerbeek, Peter Aartsma, Otar Abdalaze, Mehdi Abedi, Rien Aerts, Negar Ahmadian, Antje Ahrends, Juha M. Alatalo, Jake M Alexander, Camille Nina Allonsius, Jan Altman, Christof Ammann, Christian Andres, Christopher Andrews, Jonas Ardö, Nicola Arriga, Alberto Arzac, Valeria Aschero, Rafael L Assis, Jakob Johann Assmann, Maaike Y Bader, Khadijeh Bahalkeh, Peter Barančok, Isabel C. Barrio, Agustina Barros, Matti Barthel, Edmund W Basham, Marijn Bauters, Manuele Bazzichetto, Luca Belelli Marchesini, Michael C Bell, Juan C Benavides, José Luis Benito Alonso, Bernd J Berauer, Jarle W Bjerke, Robert G Björk, Mats P Björkman, Katrin Björnsdóttir, Benjamin Blonder, Pascal Boeckx, Julia Boike, Stef Bokhorst, Bárbara N S Brum, Josef Brůna, Nina Buchmann, Pauline Buysse, José Luís Camargo, Otávio C Campoe, Onur Candan, Rafaella Canessa, Nicoletta Cannone, Michele Carbognani, Jofre Carnicer, Angélica Casanova-Katny, Simone Cesarz, Bogdan Chojnicki, Philippe Choler, Steven L Chown, Edgar F Cifuentes, Marek Čiliak, Tamara Contador, Peter Convey, Elisabeth J Cooper, Edoardo Cremonese, Salvatore R Curasi, Robin Curtis, Maurizio Cutini, C Johan Dahlberg, Gergana N Daskalova, Miguel Angel de Pablo, Stefano Della Chiesa, Jürgen Dengler, Bart Deronde, Patrice Descombes, Valter Di Cecco, Michele Di Musciano, Jan Dick, Romina D Dimarco, Jiri Dolezal, Ellen Dorrepaal, Jiří Dušek, Nico Eisenhauer, Lars Eklundh, Todd E Erickson, Brigitta Erschbamer, Werner Eugster, Robert M Ewers, Dan A Exton, Nicolas Fanin, Fatih Fazlioglu, Iris Feigenwinter, Giuseppe Fenu, Olga Ferlian, M Rosa Fernández Calzado, Eduardo Fernández-Pascual, Manfred Finckh, Rebecca Finger Higgens, T'ai G W Forte, Erika C Freeman, Esther R Frei, Eduardo Fuentes-Lillo, Rafael A García, María B García, Charly Géron, Mana Gharun, Dany Ghosn, Khatuna Gigauri, Anne Gobin, Ignacio Goded, Mathias Goeckede, Felix Gottschall, Keith Goulding, Sanne Govaert, Bente Jessen Graae, Sarah Greenwood, Caroline Greiser, Achim Grelle, Benoit Guénard, Mauro Guglielmin, Joannès Guillemot, Peter Haase, Sylvia Haider, Aud H Halbritter, Maroof Hamid, Albin Hammerle, Arndt Hampe, Siri V Haugum, Lucia Hederová, Bernard Heinesch, Carole Helfter, Daniel Hepenstrick, Maximiliane Herberich, Mathias Herbst, Luise Hermanutz, David S. Hik, Raúl Hoffrén, Jürgen Homeier, Lukas Hörtnagl, Toke T. Høye, Filip Hrbacek, Kristoffer Hylander, Hiroki Iwata, Marcin Antoni Jackowicz-Korczynski, Hervé Jactel, Järvi Järveoja, Szymon Jastrzębowski, Anke Jentsch, Juan J Jiménez, I.S. Jonsdottir, Tommaso Jucker, Alistair S Jump, Radoslaw Juszczak, Róbert Kanka, Vít Kašpar, George Kazakis, Julia Kelly, Anzar A Khuroo, Leif Klemedtsson, Marcin Klisz, Natascha Kljun, Alexander Knohl, Johannes Kobler, Jozef Kollár, Martyna M Kotowska, Bence Kovács, Jürgen Kreyling, Andrea Lamprecht, Simone I Lang, Christian Larson, Keith Larson, Kamil Laska, Guerric le Maire, Rachel I Leihy, Luc Lens, Bengt Liljebladh, Annalea Lohila, Juan Lorite, Benjamin Loubet, Joshua Lynn, Martin Macek, Roy Mackenzie, Enzo Magliulo, Regine Maier, Francesco Malfasi, František Máliš, Matěj Man, Giovanni Manca, Antonio Manco, Tanguy Manise, Paraskevi Manolaki, Felipe Marciniak, Radim Matula, Ana Clara Mazzolari, Sergiy Medinets, Volodymyr Medinets, Camille Meeussen, Sonia Merinero, Rita de Cássia Guimarães Mesquita, Katrin Meusburger, F.J.R. Meysman, Sean T Michaletz, Ann Milbau, Dmitry Moiseev, Pavel Moiseev, Andrea Mondoni, Ruth Monfries, Leonardo Montagnani, Mikel Moriana-Armendariz, Umberto Morra di Cella, Martin Mörsdorf, Jonathan R Mosedale, Lena Muffler, Miriam Muñoz-Rojas, Jonathan A Myers, Isla H Myers-Smith, László Laszlo Nagy, Marianna Nardino, Ilona Naujokaitis-Lewis, Emily Newling, Lena Nicklas, Georg Niedrist, Armin Niessner, Mats B Nilsson, Signe Normand, Marcelo D Nosetto, Yann Nouvellon, Martin A Nuñez, Romà Ogaya, Jérôme Ogée, Joseph Okello, Janusz Olejnik, Jørgen Eivind Olesen, Øystein Opedal, Simone Orsenigo, Andrej Palaj, Timo Pampuch, Alexey V Panov, Meelis Pärtel, Ada Pastor, Aníbal Pauchard, Harald Pauli, Marian Pavelka, William D Pearse, Matthias Peichl, Loïc Pellissier, Rachel M Penczykowski, Josep Peñuelas, Matteo Petit Bon, Alessandro Petraglia, Shyam S Phartyal, Gareth K Phoenix, Casimiro Pio, Andrea Pitacco, Camille Pitteloud, Roman Plichta, Francesco Porro, Miguel Portillo-Estrada, Jérôme Poulenard, Rafael Poyatos, Anatoly S Prokushkin, Radoslaw Puchalka, Mihai Pușcaș, Dajana Radujković, Krystal Randall, Amanda Ratier Backes, Sabine Remmele, Wolfram Remmers, David Renault, Anita C. Risch, Christian Rixen, Sharon A Robinson, Bjorn J.M. Robroek, Adrian V. Rocha, Christian Rossi, Graziano Rossi, Olivier Roupsard, Alexey V Rubtsov, Patrick Saccone, Clotilde Sagot, Jhonatan Sallo Bravo, Cinthya C Santos, Judith Sarneel, Tobias Scharnweber, Jonas Schmeddes, Marius Schmidt, Thomas Scholten, Max A. Schuchardt, Naomi Schwartz, Tony Scott, Julia Seeber, Ana Cristina Segalin de Andrade, Tim Seipel, Philipp Semenchuk, Rebecca A. Senior, Josep M Serra-Diaz, Piotr Sewerniak, Ankit Shekhar, Nikita V Sidenko, Lukas Siebicke, Laura Siegwart Collier, Elizabeth Simpson, David P Siqueira, Zuzana Sitková, Johan Six, Marko Smiljanic, Stuart W Smith, Sarah Smith-Tripp, Ben Somers, Mia Vedel Sørensen, José João L L Souza, Bartolomeu Israel Souza, Arildo Souza Dias, Marko J Spasojevic, James D.M. Speed, Fabien Spicher, Angela Stanisci, Klaus Steinbauer, Rainer Steinbrecher, Michael Steinwandter, Michael Stemkovski, Jörg G. Stephan, Christian Stiegler, Stefan Stoll, Martin Svátek, Miroslav Svoboda, Torbern Tagesson, Andrew J Tanentzap, Franziska Tanneberger, Jean-Paul Theurillat, Haydn J D Thomas, Andrew D Thomas, Katja Tielbörger, Marcello Tomaselli, Urs Albert Treier, Mario Trouillier, Pavel Dan Turtureanu, Rosamond Tutton, Vilna A Tyystjärvi, Masahito Ueyama, Karol Ujházy, Mariana Ujházyová, Domas Uogintas, Anastasiya V Urban, Josef Urban, Marek Urbaniak, Tudor-Mihai Ursu, Francesco Primo Vaccari, Stijn Van de Vondel, Liesbeth van den Brink, Maarten Van Geel, Vigdis Vandvik, Pieter Vangansbeke, Andrej Varlagin, Ciska Veen, Elmar M. Veenendaal, Susanna E Venn, Hans Verbeeck, Erik Verbruggen, Frank G A Verheijen, Luis Villar, Luca Vitale, Pascal Vittoz, Maria Vives-Ingla, Jonathan von Oppen, Josefine Walz, Runxi Wang, Yifeng Wang, Robert G Way, Ronja E M Wedegärtner, Robert Weigel, Jan Wild, Matthew Wilkinson, Martin Wilmking, Lisa Wingate, Manuela Winkler, Sonja Wipf, Georg Wohlfahrt, Georgios Xenakis, Yan Yang, Zicheng Yu, Kailiang Yu, Florian Zellweger, Jian Zhang, Zhaochen Zhang, Peng Zhao, Klaudia Ziemblińska, Reiner Zimmermann, Shengwei Zong, Viacheslav I Zyryanov, Ivan Nijs, Jonathan Lenoir

    Research in global change ecology relies heavily on global climatic grids derived from estimates of air temperature in open areas at around 2 m above the ground. These climatic grids do not reflect conditions below vegetation canopies and near the ground surface, where critical ecosystem functions occur and most terrestrial species reside. Here, we provide global maps of soil temperature and bioclimatic variables at a 1-km 2 resolution for 0-5 and 5-15 cm soil depth. These maps were created by calculating the difference (i.e. offset) between in situ soil temperature measurements, based on time series from over 1200 1-km 2 pixels (summarized from 8519 unique temperature sensors) across all the world's major terrestrial biomes, and coarse-grained air temperature estimates from ERA5-Land (an atmospheric reanalysis by the European Centre for Medium-Range Weather Forecasts). We show that mean annual soil temperature differs markedly from the corresponding gridded air temperature, by up to 10°C (mean = 3.0 ± 2.1°C), with substantial variation across biomes and seasons. Over the year, soils in cold and/or dry biomes are substantially warmer (+3.6 ± 2.3°C) than gridded air temperature, whereas soils in warm and humid environments are on average slightly cooler (-0.7 ± 2.3°C). The observed substantial and biome-specific offsets emphasize that the projected impacts of climate and climate change on near-surface biodiversity and ecosystem functioning are inaccurately assessed when air rather than soil temperature is used, especially in cold environments. The global soil-related bioclimatic variables provided here are an important step forward for any application in ecology and related disciplines. Nevertheless, we highlight the need to fill remaining geographic gaps by collecting more in situ measurements of microclimate conditions to further enhance the spatiotemporal resolution of global soil temperature products for ecological applications.
  • Journal of Ecology

    Steering the soil microbiome by repeated litter addition

    Ciska Veen, Freddy ten Hooven, Carolin Weser, Emilia Hannula
    1. Microbial communities drive plant litter breakdown. Litters originating from different plant species are often associated with specialised microbiomes that accelerate the breakdown of that litter, known as home-field advantage. Yet, how and how fast microbial communities specialise towards litter inputs is not known.
    2. Here we study effects of repeated litter additions on soil microbial community structure and functioning. We set up a 9-month, full-factorial, reciprocal litter transplant experiment with soils and litters from six plant species (three grasses and three trees). We measured fungal and bacterial community composition, litter mass loss and home-field effects.
    3. We found that repeated litter additions resulted in convergence in fungal community composition driven by litter functional group (trees vs. grasses). Grasses enriched Sordariomycetes, while Tremellomycetes, Eurotiomycetes and Leotiomycetes were favoured by tree litter. Bacterial community composition, litter mass loss and home-field effects were not affected by litter incubation, but there was a relationship between fungal community composition and mass loss.
    4. We conclude that repeated litter incubations can result in directional shifts in fungal community composition, while 9 months of litter addition did not change bacterial community composition and the functioning and specialisation of microbial communities.
    5. Testing further how repeated litter inputs affect microbial functioning is essential for steering decomposer communities for optimal soil carbon and nutrient cycling.
  • ISME Journal

    Protists as catalyzers of microbial litter breakdown and carbon cycling at different temperature regimes

    Stefan Geisen, Shunran Hu, Thomas Edison E. dela Cruz, Ciska Veen
    Soil bacteria and fungi are key drivers of carbon released from soils to the atmosphere through decomposition of plant-derived organic carbon sources. This process has important consequences for the global climate. While global change factors, such as increased temperature, are known to affect bacterial- and fungal-mediated decomposition rates, the role of trophic interactions in affecting decomposition remains largely unknown. We designed synthetic microbial communities consisting of eight bacterial and eight fungal species and tested the influence of predation by a model protist, Physarum polycephalum, on litter breakdown at 17 and 21 °C. Protists increased CO2 release and litter mass loss by ~35% at 17 °C lower temperatures, while they only had minor effects on microbial-driven CO2 release and mass loss at 21 °C. We found species-specific differences in predator–prey interactions, which may affect microbial community composition and functioning and thus underlie the impact of protists on litter breakdown. Our findings suggest that microbial predation by fast-growing protists is of under-appreciated functional importance, as it affects decomposition and, as such, may influence global carbon dynamics. Our results indicate that we need to better understand the role of trophic interactions within the microbiome in controlling decomposition processes and carbon cycling.
  • Ecology

    Belowground community turnover accelerates the decomposition of standing dead wood

    Mark Bradford, Daniel S. Maynard, Tom Crowther, Paul T. Frankson, Jacqueline E. Mohan, Corinna Steinrueck, Ciska Veen, Joshua R. King, Robert J. Warren

    Standing dead trees (snags) decompose more slowly than downed dead wood and provide critical habitat for many species. The rate at which snags fall therefore influences forest carbon dynamics and biodiversity. Fall rates correlate strongly with mean annual temperature, presumably because warmer climates facilitate faster wood decomposition and hence degradation of the structural stability of standing wood. These faster decomposition rates coincide with turnover from fungal-dominated wood decomposer communities in cooler forests to codomination by fungi and termites in warmer regions. A key question for projecting forest dynamics is therefore whether temperature effects on wood decomposition arise primarily because warmer conditions facilitate faster decomposer metabolism, or are also influenced indirectly by belowground community turnover (e.g., termites exert additional influence beyond fungal-plus-bacterial mediated decomposition). To test between these possibilities, we simulate standing dead trees with untreated wooden posts and follow them in the field across 5 yr at 12 sites, before measuring buried, soil–air interface and aerial post sections to quantify wood decomposition and organism activities. High termite activities at the warmer sites are associated with rates of postfall that are three times higher than at the cooler sites. Termites primarily consume buried wood, with decomposition rates greatest where termite activities are highest. However, where higher microbial and termite activities co-occur, they appear to compensate for one another first, and then to slow decomposition rates at their highest activities, suggestive of interference competition. If the range of microbial and termite codomination of wood decomposer communities expands under climate warming, our data suggest that expansion will accelerate snag fall with consequent effects on forest carbon cycling and biodiversity in forests previously dominated by microbial decomposers.
  • Journal of Plant Ecology

    The role of soil-borne fungi in driving the coexistence of Pinus massoniana and Lithocarpus glaber in a subtropical forest via plant-soil feedback

    Yumei Pan, Zhongyuan Yao , Naili Zhang, Ciska Veen
    Aims: Plant-soil feedback (PSF) is a key mechanism that can facilitate tree species coexistence and diversity. Substantial evidence suggests that species-specific soil-borne pathogens around adult trees limit the performance of home (conspecific) seedlings relative to foreign (heterospecific) seedlings. However, the underlying mechanism remains largely elusive. Methods: Here, we conducted a reciprocal transplant pot experiment using seedlings and from two tree species, Pinus massoniana and Lithocarpus glaber that are dominant and coexist in a subtropical, evergreen, broad-leaf forest in Gutianshan, Zhejiang Province of eastern China. We examined how seedlings from the two tree species responded to soils originating from underneath their own versus the other tree species, using a full-factorial design. Additionally, we added a fungicide (benomyl) to half of the pots to evaluate the role of soil-borne fungi on seedling growth. Important Findings: We found that the seedlings from L. glaber grew better in soils that were collected from beneath the canopy of P. massoniana, while seedling growth of P. massioniana was not affected by soil origin. The addition of fungicide benomyl resulted in a shift towards more positive PSF effects for L. glaber, indicating that L. glaber seedlings performed better in their own soils than in soils from P. massoniana in the absence of fungi. Our findings highlight the importance of soil-borne pathogenic and ectomycorrhizal fungi in driving PSF, and indicate that PSF may promote the coexistence of two subtropical tree species by reducing the performance of L. glaber in own soils.
  • Trends in Ecology and Evolution

    Plant–Soil Feedbacks and Temporal Dynamics of Plant Diversity–Productivity Relationships

    Maddy Thakur, Wim H. van der Putten, Rutger Wilschut, Ciska Veen, Paul Kardol, Jasper van Ruijven, Eric Allan, Christiane Roscher, Mark van Kleunen, T. Martijn Bezemer

    Plant–soil feedback (PSF) and diversity–productivity relationships are important research fields to study drivers and consequences of changes in plant biodiversity. While studies suggest that positive plant diversity–productivity relationships can be explained by variation in PSF in diverse plant communities, key questions on their temporal relationships remain. Here, we discuss three processes that change PSF over time in diverse plant communities, and their effects on temporal dynamics of diversity–productivity relationships: spatial redistribution and changes in dominance of plant species; phenotypic shifts in plant traits; and dilution of soil pathogens and increase in soil mutualists. Disentangling these processes in plant diversity experiments will yield new insights into how plant diversity–productivity relationships change over time.
  • Soil Biology and Biochemistry

    Optimizing stand density for climate-smart forestry: a way forward towards resilient forests with enhanced carbon storage under extreme climate events

    Frank Sterck, Marleen Vos, Emilia Hannula, Steven de Goede, Wim de Vries, Jan den Ouden, Gert-Jan Nabuurs, Wim H. van der Putten, Ciska Veen
    As a response to the increased pressure of global climate change on most ecosystems, national and international agreements aim at creating forests that are productive, resilient to climate change, and that store carbon to mitigate global warming. However, these aims are being challenged by increased tree mortality rates and decreased tree growth rates in response to increased incidence of extreme drought events. These phenomena make us aware of a lack of crucial insights into the effects of forest management on the growth and survival of trees, and on carbon storage in both trees and forest soils under increased incidence of drought. Here we compile current knowledge on how forest management and drought impact on tree growth and survival, and above- and belowground carbon storage in forest ecosystems. Based on this, we propose that climate-smart forestry may benefit from controlling stand density at intermediate levels (>60%, e.g.∼80%) by applying low levels of tree harvest intensity on a regular base. Furthermore, we propose that the actual optimal density will depend on the tree species, site conditions and management history. As a next step, studies are needed that take an above- and belowground approach and combine forest experiments with mechanistic models on water, carbon and nutrient flows in trees and soils within forests in order to transform current results, which focus on either soil or trees and are often highly-context dependent, to a more generic forest framework. Such a generic framework would be needed to enhance understanding across forest ecosystems on how forest management may promote forest resilience, productivity and carbon storage with increasing drought.
  • New Phytologist

    Home-field advantage of litter decomposition

    Nicolas Fanin, D. Lin, G.T. Freschet, A.D. Keiser, L. Augusto, David A. Wardle, Ciska Veen
    Plants often associate with specialized decomposer communities that increase plant litter breakdown, a phenomenon that is known as the ‘home-field advantage’ (HFA). Although the concept of HFA has long considered only the role of the soil microbial community, explicit consideration of the role of the microbial community on the foliage before litter fall (i.e. the phyllosphere community) may help us to better understand HFA. We investigated the occurrence of HFA in the presence vs absence of phyllosphere communities and found that HFA effects were smaller when phyllosphere communities were removed. We propose that priority effects and interactions between phyllosphere and soil organisms can help explain the positive effects of the phyllosphere at home, and suggest a path forward for further investigation.
  • PLoS One

    The abundance of arbuscular mycorrhiza in soils is linked to the total length of roots colonized at ecosystem level

    M Barcelo, P.M. Van Bodegom, L. Tedersoo, N. den Haan, Ciska Veen, I Ostonen, K. B. Trimbos, Nadia A. Soudzilovskaia
    Arbuscular mycorrhizal fungi (AMF) strongly affect ecosystem functioning. To understand and quantify the mechanisms of this control, knowledge about the relationship between the actual abundance and community composition of AMF in the soil and in plant roots is needed. We collected soil and root samples in a natural dune grassland to test whether, across a plant community, the abundance of AMF in host roots (measured as the total length of roots colonized) is related to soil AMF abundance (using the neutral lipid fatty acids (NLFA) 16:1ω5 as proxy). Next-generation sequencing was used to explore the role of community composition in abundance patterns. We found a strong positive relationship between the total length of roots colonized by AMF and the amount of NLFA 16:1ω5 in the soil. We provide the first field-based evidence of proportional biomass allocation between intra-and extraradical AMF mycelium, at ecosystem level. We suggest that this phenomenon is made possible by compensatory colonization strategies of individual fungal species. Finally, our findings open the possibility of using AMF total root colonization as a proxy for soil AMF abundances, aiding further exploration of the AMF impacts on ecosystems functioning.
  • Global Change Biology

    Nutrient availability controls the impact of mammalian herbivores on soil carbon and nitrogen pools in grasslands

    Judith Sitters, Jasper Wubs, (Liesbeth) E.S. Bakker, Tom Crowther, Peter B. Adler, Sumanta Bagchi, Jonathan D. Bakker, L. Biederman, Elizabeth T. Borer, Elsa E. Cleland, Nico Eisenhauer, J Firn, Laureano Gherardi, Nicole Hagenah, Yann Hautier, Sarah Hobbie, Johannes M. H. Knops, A.S. MacDougall, Rebecca L. McCulley, Joslin L. Moore, Brent Mortensen, Pablo L. Peri, Suzanne M. Prober, C Riggs, Anita C. Risch, Martin Schütz, Eric W. Seabloom, Julia Siebert, Carly J. Stevens, Ciska Veen
    Grasslands are subject to considerable alteration due to human activities globally, including widespread changes in populations and composition of large mammalian herbivores and elevated supply of nutrients. Grassland soils remain important reservoirs of carbon (C) and nitrogen (N). Herbivores may affect both C and N pools and these changes likely interact with increases in soil nutrient availability. Given the scale of grassland soil fluxes, such changes can have striking consequences for atmospheric C concentrations and the climate. Here, we use the Nutrient Network experiment to examine the responses of soil C and N pools to mammalian herbivore exclusion across 22 grasslands, under ambient and elevated nutrient availabilities (fertilized with NPK + micronutrients). We show that the impact of herbivore exclusion on soil C and N pools depends on fertilization. Under ambient nutrient conditions, we observed no effect of herbivore exclusion, but under elevated nutrient supply, pools are smaller upon herbivore exclusion. The highest mean soil C and N pools were found in grazed and fertilized plots. The decrease in soil C and N upon herbivore exclusion in combination with fertilization correlated with a decrease in aboveground plant biomass and microbial activity, indicating a reduced storage of organic matter and microbial residues as soil C and N. The response of soil C and N pools to herbivore exclusion was contingent on temperature – herbivores likely cause losses of C and N in colder sites and increases in warmer sites. Additionally, grasslands that contain mammalian herbivores have the potential to sequester more N under increased temperature variability and nutrient enrichment than ungrazed grasslands. Our study highlights the importance of conserving mammalian herbivore populations in grasslands worldwide. We need to incorporate local‐scale herbivory, and its interaction with nutrient enrichment and climate, within global‐scale models to better predict land–atmosphere interactions under future climate change.
  • One Earth

    Climate Extremes, Rewilding, and the Role of Microhabitats

    Maddy Thakur, (Liesbeth) E.S. Bakker, Ciska Veen, Jeff A. Harvey

    Climate extremes are expected to become more commonplace and more severe, putting species and ecosystems at unprecedented risks. We recommend that rewilding programs can create conditions for ecosystems to endure and recover rapidly from climate extremes by incorporating ecosystem engineers of various body sizes and life forms.
  • Global Ecology and Biogeography

    Nonlinear responses of soil nematode community composition to increasing aridity

    Dan Xiong, Cunzheng Wei, Jasper Wubs, Ciska Veen, W. Liang, X. Wang, L. Qi, Wim H. van der Putten, Xingguo Han
  • Oikos

    Herbivore phenology can predict response to changes in plant quality by livestock grazing

    Yu Zhu, Ciska Veen, Deli Wang, Ling Wang, Zhiwei Zhong, Quanhui Ma, Heng Li, Xincheng Li, Duofeng Pan, (Liesbeth) E.S. Bakker
    Livestock grazing can have a strong impact on herbivore abundance, distribution and community. However, not all species of herbivores respond the same way to livestock grazing, and we still have a poor understanding of the underlying mechanisms driving these differential responses. Here, we investigate the effect of light intensity cattle grazing on the abundance of two grasshoppers (Euchorthippus cheui and E. unicolor) that co-occur in the same grasslands and feed on the same food plant (the dominant grass Leymus chinensis). The two grasshopper species differ in phenology so that their peak abundances are separated into early- and late-growing seasons. We used an exclosure experiment to monitor grasshopper abundance and food quality in the field under grazed and ungrazed conditions, and performed feeding trials to examine grasshopper preference for grazed or ungrazed food plants in the laboratory. We found that the nitrogen content of L. chinensis leaves continuously declined in the ungrazed areas, but was significantly enhanced by cattle grazing over the growing season. Cattle grazing facilitated the early-season grasshopper E. cheui, whereas it suppressed the late-season grasshopper E. unicolor. Moreover, feeding trials showed that E. cheui preferred L. chinensis from grazed plots, while E. unicolor preferred the leaves from ungrazed plots. We conclude that livestock grazing has opposite effects on the two grasshopper species, and that these effects may be driven by grazing-induced changes in plant nutrient content and the unique nutritional niches of the grasshoppers. These results suggest that insects that belong to the same guild can have opposite nutrient requirements, related to their distinct phenologies, and that this can ultimately affect their response to cattle grazing. Our results show that phenology may link insect physiological needs to local resource availabilities, and should be given more attention in future work on interactions between large herbivores and insects.
  • Soil Biology & Biochemistry

    Short-term temperature history affects mineralization of fresh litter and extant soil organic matter, irrespective of agricultural management

    Kyle Mason-Jones, Pim Vrehen, Kevin Koper, Jin Wang, Wim H. van der Putten, Ciska Veen
    The influence of temperature on mineralization of plant litter and pre-existing soil organic matter (SOM) involves not only the prevailing temperature, but also how it has changed through time. However, little is known about how temperature variability through time influences mineralization processes. Here, we investigated how short-term temperature history affects the mineralization of SOM and plant litter in soils from different agricultural management systems. We used soils from a long-term experiment with conventional and organic management treatments to set up microcosms. The microcosms were exposed to eight days of contrasting temperature regimes (different mean temperatures and constant versus fluctuating temperatures). Microcosms were then returned to a common temperature of 16 °C, 13C-labelled plant litter was added to half of them, and CO2 efflux was measured over the following week. We found that SOM and litter mineralization were both sensitive to the temperature history, with lower mean temperatures during preliminary treatment associated with higher mineralization during the subsequent common-temperature incubation. This effect persisted through the week after temperature differences were removed. Different patterns of temperature fluctuation and agricultural management did not significantly affect mineralization during common-temperature incubation. The history sensitivity of litter mineralization, despite litter being added after temperature differences had ended, indicates that the temperature history effects may be driven by short-term microbial acclimation. We conclude that organic matter and litter mineralization, which are key processes in the carbon cycle, are sensitive to short-term temperature history. This suggests that future investigations of soil CO2 efflux may need to take recent weather effects into account.
  • Soil Biology & Biochemistry

    ‘Home’ and ‘away’ litter decomposition depends on the size fractions of the soil biotic community

    Yingbin Li, Ciska Veen, (Gera) W.H.G. Hol, Simon Vandenbrande, Emilia Hannula, Freddy ten Hooven, Qi Li, Wenju Liang, T. Martijn Bezemer
    The ‘home-field advantage’ (HFA) hypothesis predicts that litter decomposition is accelerated in its home environment (i.e. in conspecific soil). Soil organisms play a key role in driving such HFA effects. Soil biota have a large range of body sizes, referred to as size fractions, which may influence their roles in the decomposition process and in the generation of HFA effects. However, how HFA effects depend on the different size fractions of the soil biotic community is unknown. We conducted a microcosm decomposition experiment to examine how size fractions of the soil biotic community affected litter decomposition and HFA effects. In a semi-natural grassland in the Netherlands, we collected leaf litter and soil from two abundant forbs: Tanacetum vulgare and Jacobaea vulgaris. Watery extracts of the soils were sieved through differently-meshed sieves (ranging from 850 μm to 6 μm) to obtain soil communities of different size fractions. Microcosms were inoculated with these different size fractions of the soil biotic community and we examined their effects on microbial composition, litter mass loss and HFA effects. Three months after inoculation, the diversity of the fungal community in the inoculated pots decreased with decreasing size fractions of the soil biotic community. Similarly, litter mass loss also decreased with decreasing soil biotic community size. In contrast, the HFA effect increased with decreasing size fractions of the soil biotic community, but these differences disappeared after six months of decomposition. Our results indicate that soil microorganisms, mainly the smallest size fractions, are specialized to decompose specific resources and thus promote HFA effects, but that their effect is only apparent during specific stages of litter decomposition.
  • Global Ecology and Biogeography

    Soil microbial biomass increases along elevational gradients in the tropics and subtropics but not elsewhere

    X. He, E. Hou, Ciska Veen, M. Ellwood, Paul Dijkstra, X Sui, S. Zhang, D Wen, C Chu
  • Frontiers in Microbiology

    Belowground Consequences of Intracontinental Range-Expanding Plants and Related Natives in Novel Environments

    Marta Manrubia-Freixa, Basten Snoek, Carolin Weser, Ciska Veen, Wim H. van der Putten
    Introduced exotic plant species that originate from other continents are known to alter soil microbial community composition and nutrient cycling. Plant species that expand range to higher latitudes and altitudes as a consequence of current climate warming might as well affect the composition and functioning of native soil communities in their new range. However, the functional consequences of plant origin have been poorly studied in the case of plant range shifts. Here, we determined rhizosphere bacterial communities of four intracontinental range-expanding plant species in comparison with their four congeneric natives grown in soils collected from underneath those plant species in the field and in soils that are novel to them. We show that, when controlling for both species relatedness and soil characteristics, range-expanding plant species in higher latitude ecosystems will influence soil bacterial community composition and nutrient cycling in a manner similar to congeneric related native species. Our results highlight the importance to include phylogenetically controlled comparisons to disentangle the effect of origin from the effect of contrasting plant traits in the context of exotic plant species.
  • Functional Ecology

    Soil functional responses to drought under range-expanding and native plant communities

    Marta Manrubia-Freixa, Wim H. van der Putten, Carolin Weser, Freddy ten Hooven, Henk Martens, Pella Brinkman, Stefan Geisen, Kelly Ramirez, Ciska Veen
    Current climate warming enables plant species and soil organisms to expand their range to higher latitudes and altitudes. At the same time, climate change increases the incidence of extreme weather events such as drought. While it is expected that plants and soil organisms originating from the south are better able to cope with drought, little is known about the consequences of their range shifts on soil functioning under drought events.

    Here, we test how range‐expanding plant species and soil communities may influence soil functioning under drought. We performed a full‐factorial outdoor mesocosm experiment with plant communities of range expanders or related natives, with soil inocula from the novel or the original range, with or without summer drought. We measured litter decomposition, carbon mineralization and enzyme activities, substrate‐induced respiration, and the relative abundance of soil saprophytic fungi immediately after drought and at 6 and 12 weeks after rewetting.

    Drought decreased all soil functions regardless of plant and soil origin except one; soil respiration was less reduced in soils of range‐expanding plant communities, suggesting stronger resistance to drought. After rewetting, soil functioning responses depended on plant and soil origin. Soils of native plant communities with a history of drought had more litter mass loss and higher relative abundance of saprophytic fungi than soils without drought and soils of range expanders. Functions of soil from range expanders recovered in a more conservative manner than soils of natives, as litter mass loss did not exceed the control rates. At the end of the experiment, after rewetting, most soil functions in mesocosms with drought history did not differ anymore from the control.

    We conclude that functional consequences of range expanding plants and soil biota may interact with effects of drought, and that these effects are most prominent during the first weeks after rewetting of the soil.
  • Frontiers in Ecology and Evolution

    Applying the aboveground-belowground interaction concept in agriculture

    Ciska Veen, Jasper Wubs, Richard D. Bardgett, Edmundo Barrios, Mark Bradford, Sabrina Almeida de Carvalho, Gerlinde De Deyn, Franciska T. De Vries, Ken E. Giller, David Kleijn, Douglas A. Landis, Walter A.H. Rossing, Maarten Schrama, Johan Six, Paul C. Struik, Stijn van Gils, Johannes S.C. Wiskerke, Wim H. van der Putten, Louise E.M. Vet

    Interactions between aboveground and belowground organisms are important drivers of plant growth and performance in natural ecosystems. Making practical use of such above-belowground biotic interactions offers important opportunities for enhancing the sustainability of agriculture, as it could favor crop growth, nutrient supply, and defense against biotic and abiotic stresses. However, the operation of above-and belowground organisms at different spatial and temporal scales provides important challenges for application in agriculture. Aboveground organisms, such as herbivores and pollinators, operate at spatial scales that exceed individual fields and are highly variable in abundance within growing seasons. In contrast, pathogenic, symbiotic, and decomposer soil biota operate at more localized spatial scales from individual plants to patches of square meters, however, they generate legacy effects on plant performance that may last from single to multiple years. The challenge is to promote pollinators and suppress pests at the landscape and field scale, while creating positive legacy effects of local plant-soil interactions for next generations of plants. Here, we explore the possibilities to improve utilization of above-belowground interactions in agro-ecosystems by considering spatio-temporal scales at which aboveground and belowground organisms operate. We identified that successful integration of above-belowground biotic interactions initially requires developing crop rotations and intercropping systems that create positive local soil legacy effects for neighboring as well subsequent crops. These configurations may then be used as building blocks to design landscapes that accommodate beneficial aboveground communities with respect to their required resources. For successful adoption of above-belowground interactions in agriculture there is a need for context-specific solutions, as well as sound socio-economic embedding.
  • Functional Ecology

    Relationships between fungal community composition in decomposing leaf litter and home-field advantage effects

    Ciska Veen, Basten Snoek, Tanja Bakx-Schotman, David A. Wardle, Wim H. van der Putten
    Increasing evidence suggests that specific interactions between microbial decomposers and plant litter, named home field advantage (HFA), influence litter breakdown. However, we still have limited understanding of whether HFA relates to specific microbiota, and whether specialized microbes originate from the soil or from the leaf microbiome. Here, we disentangle the roles of soil origin, litter types, and the microbial community already present on the leaf litter in determining fungal community composition on decomposing leaf litter and HFA.

    We collected litters and associated soil samples from a secondary succession gradient ranging from herbaceous vegetation on recently abandoned ex‐arable fields to forest representing the end stage of succession. In a greenhouse, sterilized and unsterilized leaf litters were decomposed for 12 months in soils from early to late successional stages according to a full factorial design. At the end, we examined fungal community composition on the decomposing litter.

    Fungal communities on decomposed late‐successional litter in late‐successional soil differed from those in early‐ and mid‐successional stage litter and soil combinations. Soil source had the strongest impact on litter fungal composition when using sterilized litter, while the impact of litter type was strongest when using unsterilized litter. Overall, we observed HFA, as litter decomposition was accelerated in home soils. Increasing HFA did not relate to the dissimilarity in overall fungal composition, but there was increasing dissimilarity in the relative abundance of the most dominant fungal taxon between decomposing litter in home and away soils.

    We conclude that early, mid and late succession litter types did not exert strong selection effects on colonization by microorganisms from the soil species pool. Instead, fungal community composition on decomposing litter differed substantially between litter types for unsterilized litter, suggesting that the leaf microbiome, either directly or indirectly, is an important determinant of fungal community composition on decomposing leaves. HFA related most strongly to the abundance of the most dominant fungal taxa on the decomposing litter, suggesting that HFA may be attributed to some specific dominant fungi rather than to responses of the whole fungal community.
  • Functional Ecology

    Why are plant-soil feedbacks so unpredictable, and what to do about it?

    Jon De Long, Ellen L. Fry, Ciska Veen, Paul Kardol
    1.The study of feedbacks between plants and soils (plant‐soil feedbacks; PSFs) is receiving increased attention. However, PSFs have been mostly studied in isolation of abiotic and biotic drivers that could affect their strength and direction. This is problematic because it has led to limited predictive power of PSFs in ‘the real world’, leaving large knowledge gaps in our ability to predict how PSFs contribute to ecosystem processes and functions.

    2.Here, we present a synthetic framework to elucidate how abiotic and biotic drivers affect PSFs. We focus on two key abiotic drivers (temperature and soil moisture) and two key biotic drivers (aboveground plant consumers and belowground top‐down control of pathogens and mutualists). We focus on these factors because they are known drivers of plants and soil organisms and the ecosystem processes they control, and hence would be expected to strongly influence PSFs.

    3.Our framework describes the proposed mechanisms behind these drivers and explores their effects on PSFs. We demonstrate the impacts of these drivers using the fast‐ to slow‐growing plant economics spectrum. We use this well‐established paradigm because plants on opposite ends of this spectrum differ in their relationships with soil biota and have developed contrasting strategies to cope with abiotic and biotic environmental conditions.

    4.Finally, we present suggestions for improved experimental designs and scientific inference that will capture and elucidate the influence of above‐ and belowground drivers on PSFs. By establishing the role of abiotic and biotic drivers of PSFs, we will be able to make more robust predictions of how PSFs impact on ecosystem function.
  • Frontiers in Plant Science

    High grazing pressure of geese threatens conservation and restoration of reed belts

    (Liesbeth) E.S. Bakker, Ciska Veen, G. Ter Heerdt, Naomi Huig, Judith Sarneel
    Reed (Phragmites australis (Cav.) Trin. ex Steud.) beds are important habitat for marsh birds, but are declining throughout Europe. Increasing numbers of the native marsh bird, the Greylag goose (Anser anser L.), are hypothesized to cause reed bed decline and inhibit restoration of reed beds, but data are largely lacking. In this study, we experimentally tested the effect of grazing by Greylag geese on the growth and expansion of reed growing in belts along lake shorelines. After 5 years of protecting reed from grazing with exclosures, reed stems were over 4-fold denser and taller than in the grazed plots. Grazing pressure was intense with 50–100% of the stems being grazed among years in the control plots open to grazing. After 5 years of protection we opened half of the exclosures and the geese immediately grazed almost 100% of the reed stems. Whereas this did not affect the reed stem density, the stem height was strongly reduced and similar to permanently grazed reed. The next year geese were actively chased away by management from mid-March to mid-June, which changed the maximum amount of geese from over 2300 to less than 50. As a result, reed stem density and height increased and the reed belt had recovered over the full 6 m length of the experimental plots. Lastly, we introduced reed plants in an adjacent lake where no reed was growing and geese did visit this area. After two years, the density of the planted reed was six to nine-fold higher and significantly taller in exclosures compared to control plots where geese had access to the reed plants. We conclude that there is a conservation dilemma regarding how to preserve and restore reed belts in the presence of high densities of Greylag geese as conservation of both reed belts and high goose numbers seems infeasible. We suggest that there are three possible solutions for this dilemma: (1) effects of the geese can be mediated by goose population management, (2) the robustness of the reed marshes can be increased, and (3) at the landscape level, spatial planning can be used to configure landscapes with large reed bed reserves surrounded by unmown, unfertilized meadows.
  • Trends in Ecology and Evolution

    Plant-Soil Feedback: Bridging Natural and Agricultural Sciences

    P. Mariotte, Z. Mehrabi, T. Martijn Bezemer, Gerlinde De Deyn, A. Kulmatiski, Barbara Drigo, Marcel G. A. van der Heijden, Ciska Veen, Paul Kardol
    PSF has been extensively studied in both agricultural and natural systems, with increased activity in recent years, but a framework for integrating the concepts and principles developed in these systems is lacking.

    Interactions between soil biota and plant leaf and root traits have become an important tool in understanding PSF in wild plants, but this understanding has not yet been utilized in agricultural crop rotations.

    Soil inoculations with microbial strains are increasingly being used for steering the soil microbiome in agriculture but might also offer a promising method of restoration of degraded systems, and for controlling the spread of invasive species.

    Increasing evidence shows that PSF can play important roles in mediating ecosystem responses to forecasted climate change and extreme weather events.

    In agricultural and natural systems researchers have demonstrated large effects of plant–soil feedback (PSF) on plant growth. However, the concepts and approaches used in these two types of systems have developed, for the most part, independently. Here, we present a conceptual framework that integrates knowledge and approaches from these two contrasting systems. We use this integrated framework to demonstrate (i) how knowledge from complex natural systems can be used to increase agricultural resource-use efficiency and productivity and (ii) how research in agricultural systems can be used to test hypotheses and approaches developed in natural systems. Using this framework, we discuss avenues for new research toward an ecologically sustainable and climate-smart future.
  • Ecology

    Biodiversity-ecosystem functioning relationships in a long-term non-weeded field experiment

    Ciska Veen, Wim H. van der Putten, T. Martijn Bezemer
    Many grassland biodiversity experiments show a positive relationship between biodiversity and ecosystem functioning, however, in most of these experiments plant communities are established by sowing and natural colonization is prevented by selective weeding of non‐sown species. During ecosystem restoration, for example on abandoned fields, plant communities start on bare soil, and diversity is often manipulated in a single sowing event. How such initial plant diversity manipulations influence plant biodiversity development and ecosystem functioning is not well understood. We examined how relationships between taxonomic and functional diversity, biomass production and stability develop over 16 yr in non‐weeded plots sown with 15 species, four species, or that were not sown. We found that sown plant communities become functionally similar to unsown, naturally colonized plant communities. However, initial sowing treatments had long‐lasting effects on species composition and taxonomic diversity. We found only few relationships between biomass production, or stability in biomass production, and functional or taxonomic diversity, and the ones we observed were negative. In addition, the cover of dominant plant species was positively related to biomass production and stability. We conclude that effects of introducing plant species at the start of secondary succession can persist for a long time, and that in secondary succession communities with natural plant species dynamics diversity–functioning relationships can be weak or negative. Moreover, our findings indicate that in systems where natural colonization of species is allowed effects of plant dominance may underlie diversity–functioning relationships.
  • Functional Ecology

    Variation in home-field advantage and ability in leaf litter decomposition across successional gradients

    Ciska Veen, A.D. Keiser, Wim H. van der Putten, David A. Wardle
    It is increasingly recognized that interactions between plants and soil (a)biotic conditions can influence local decomposition processes. For example, decomposer communities may become specialized in breaking down litter of plant species that they are associated with, resulting in accelerated decomposition, known as “home‐field advantage” (HFA). Also, soils can vary inherently in their capacity to degrade organic compounds, known as “ability.” However, we have a poor understanding how environmental conditions drive the occurrence of HFA and ability.
    Here, we studied how HFA and ability change across three types of successional gradients: coastal sand dunes (primary succession), inland drift sands (primary succession) and ex‐arable fields (secondary succession). Across these gradients, litter quality (i.e. nutrient, carbon and lignin contents) increases with successional time for coastal dunes and decreases for the other two gradients.
    We performed a 12‐months reciprocal litter transplant experiment under greenhouse conditions using soils and litters collected from early‐, mid‐ and late‐successional stages of each gradient.
    We found that HFA and ability did not consistently shift with successional stage for all gradients, but were instead specific for each type of successional gradient. In coastal dunes, HFA was positive for early‐successional litter, in drift, sands it was negative for mid‐successional litter, and for ex‐arable fields, HFA increased with successional time. Ability of decomposer communities was highest in mid‐successional stages for coastal dunes and drift sands, but for ex‐arable fields, ability decreased throughout with successional time. High HFA was related to high litter C content and soil and organic matter content in soils and to low litter and soil nutrient concentrations. Ability did not consistently occur in successional stages with high or low litter quality.
    Synthesis. Our findings show that specific environmental conditions, such as changes in litter or soil quality, along environmental gradients can shape the influence of HFA and ability on decomposition. In sites with strong HFA or ability, interactions between plants, litter and decomposer communities will be important drivers of nutrient cycling and hence have the potential to feedback to plant growth.
  • Soil Biology & Biochemistry

    Relationship between home-field advantage of litter decomposition and priming of soil organic matter

    Paolo Di Lonardo, Marta Manrubia-Freixa, Wietse de Boer, Hans Zweers, Ciska Veen, Annemieke van der Wal
  • Pedobiologia

    Contrasting responses of springtails and mites to elevation and vegetation type in the sub-Arctic

    Stef Bokhorst, Ciska Veen, Maja K. Sundqvist, Jon De Long, Paul Kardol, David A. Wardle
    Climate change is affecting the species composition and functioning of Arctic and sub-Arctic plant and soil communities. Here we studied patterns in soil microarthropod (springtails and mites) communities across a gradient of increasing elevation that spanned 450 m, across which mean temperature declined by approximately 2.5 °C, in sub-Arctic Sweden. Across this gradient we characterized microarthropod communities in each of two types of vegetation, i.e., heath and meadow, to determine whether their responses to declining temperature differed with vegetation type. Mite abundance declined with increasing elevation, while springtail abundance showed the opposite response. Springtail communities were dominated by larger species at higher elevation. Mite abundance was unaffected by vegetation type, while springtail abundance was 53% higher in the heath than meadow vegetation across the gradient. Springtails but not mites responded differently to elevation in heath and meadow vegetation; hemi-edaphic species dominated in the heath at higher elevation while epi-edaphic species dominated in the meadow. Our results suggest that sub-Arctic mite and springtail communities will likely respond in contrasting ways to changes in vegetation and soil properties resulting from climate warming.
  • Functional Ecology

    Negative effects of litter richness on root decomposition in the presence of detritivores

    Y. Li, Xu Chen, Ciska Veen, Nico Eisenhauer, Yu Liang, X. Zhou, Naili Zhang, K. Ma
    1.Decomposition is a vital process underlying many ecosystem functions. Although a growing number of studies have tested how litter richness affects the decomposition of aboveground plant organs, knowledge remains limited about the decomposition of root mixtures. Here, we used a field experiment in a subtropical forest to investigate how species richness in root litter mixtures (air-dried fresh fine roots) affects the decomposition of root litter material.

    2.Based on the concept of resource complementarity, we hypothesized that root litter would decompose faster as the richness of the root litter mixture increased. In addition, we expected the presence of detritivores to modify the effect of root richness on mass loss, because detritivores might experience bottom-up effects from specific plant species and might affect microbial decomposer communities.

    3.We found that the richness level of root litter mixtures did not affect mass loss in the absence of detritivores. In the presence of detritivores, all root litter types decomposed faster. Notably, the positive effect of detritivores was stronger at low root litter richness than at high root litter richness, particularly during the early stages of decomposition (the first two sampling points) when litter mass loss was roughly double at low root litter richness compared to that at high root litter richness. The composition of the root fungal community measured at the last sampling point did not differ significantly across root richness levels, and was not affected by the presence of detritivores.

    4.Synthesis. Our findings demonstrate that detritivores modify the relationship between root litter diversity and root litter decomposition in subtropical forest ecosystems. This highlights an importance of cascade effects between different trophic organisms on ecosystem functioning.
  • Aquatic Botany

    Effects of temperature, moisture and soil type on seedling emergence and mortality of riparian plant species

    Gerard N.J. Ter Heerdt, Ciska Veen, Wim H. van der Putten, Jan P. Bakker
    Abstract Restoration of riparian plant communities on bare soil requires germination of seeds and establishment of seedlings. However, species that are present in the soil seed bank do not always establish in the vegetation. Temperature, moisture conditions and soil type could play a major role in the establishment of riparian plant communities, through impacting seedling emergence. We studied the effects of temperature, combinations of temperature and moisture conditions, and soil type on seedling emergence and mortality of perennial reeds (Typha latifolia and Phragmites australis) and annual or biannual pioneer species (Senecio congestus, Rumex maritimus and Chenopodium rubrum). The responses to the environmental conditions were species-specific and resulted in context-dependent differences in proportions of species emerging from the soil seed bank. Typha latifolia and S. congestus preferred wet or very wet conditions, C. rubrum and R. maritimus preferred dry to very dry conditions. Phragmites australis was able to establish under all conditions. Both cold and very dry conditions resulted in low emergence and survival, which was not fully compensated for when conditions became favorable again. Senecio congestus, R. maritimus and C. rubrum benefitted from secondary seedling emergence when, after a very dry period, the weather became very wet again, while T. latifolia and P. australis remained absent. When the conditions remained wet, more seedlings emerged from sand than from clay. However, when the soil was drying out, fewer seedlings emerged from sand than from clay. We propose that using information on plant species-specific responses to abiotic environmental conditions during germination, emergence and establishment can help to restore different target riparian plant communities.
  • Plant and Soil

    Legacy effects of altered flooding regimes on decomposition in a boreal floodplain

    Judith Sarneel, Ciska Veen

    Background and aims

    Since long-term experiments are scarce, we have poor understanding of how changed flooding regimes affect processes such as litter decomposition.


    We simulated short- and long-term changed flooding regimes by transplanting turfs between low (frequently flooded) and high (in-frequently flooded) elevations on the river bank in 2000 (old turfs) and 2014 (young turfs). We tested how incubation elevation, turf origin and turf age affected decomposition of standard litter (tea) and four types of local litter.


    For tea, we found that the initial decomposition rate (k) and stabilization (S) of labile material during the second decomposition phase were highest at high incubation elevation. We found intermediate values for k and S in young transplanted turfs, but turf origin was not important in old turfs. Local litter mass loss was generally highest at high incubation elevations, and effects of turf origin and turf age were litter-specific.


    We conclude that incubation elevation, i.e., the current flooding regime, was the most important factor driving decomposition. Soil origin (flooding history) affected decomposition of tea only in young turfs. Therefore, we expect that changes in flooding regimes predominantly affect decomposition directly, while indirect legacy effects are weaker and litter- or site-specific.
  • Frontiers of Earth Science

    The stoichiometry of nutrient release by terrestrial herbivores and its ecosystem consequences.

    Judith Sitters, (Liesbeth) E.S. Bakker, M. Veldhuis, Ciska Veen, Harry Olde Venterink, M.J. Vanni
    It is widely recognized that the release of nutrients by herbivores via their waste products strongly impacts nutrient availability for autotrophs. The ratios of nitrogen (N) and phosphorus (P) recycled through herbivore release (i.e., waste N:P) are mainly determined by the stoichiometric composition of the herbivore's food (food N:P) and its body nutrient content (body N:P). Waste N:P can in turn impact autotroph nutrient limitation and productivity. Herbivore-driven nutrient recycling based on stoichiometric principles is dominated by theoretical and experimental research in freshwater systems, in particular interactions between algae and invertebrate herbivores. In terrestrial ecosystems, the impact of herbivores on nutrient cycling and availability is often limited to studying carbon (C):N and C:P ratios, while the role of terrestrial herbivores in mediating N:P ratios is also likely to influence herbivore-driven nutrient recycling. In this review, we use rules and predictions on the stoichiometry of nutrient release originating from algal-based aquatic systems to identify the factors that determine the stoichiometry of nutrient release by herbivores. We then explore how these rules can be used to understand the stoichiometry of nutrient release by terrestrial herbivores, ranging from invertebrates to mammals, and its impact on plant nutrient limitation and productivity. Future studies should focus on measuring both N and P when investigating herbivore-driven nutrient recycling in terrestrial ecosystems, while also taking the form of waste product (urine or feces) and other pathways by which herbivores change nutrients into account, to be able to quantify the impact of waste stoichiometry on plant communities.
  • Nature Ecology and Evolution

    A test of the hierarchical model of litter decomposition

    Mark Bradford, Ciska Veen, A. Bonis, E.M. Bradford, Aimée T. Classen, J.H.C. Cornelissen, Tom Crowther, J.R. De Long, G.T. Freschet, Paul Kardol, Marta Manrubia-Freixa, Daniel S. Maynard, G.S. Newman, R. Van Logtestijn, Maria Viketoft, David A. Wardle, W.R. Wieder, S.A. Wood, Wim H. van der Putten
    Our basic understanding of plant litter decomposition informs the assumptions underlying widely applied soil biogeochemical models, including those embedded in Earth system models. Confidence in projected carbon cycle–climate feedbacks therefore depends on accurate knowledge about the controls regulating the rate at which plant biomass is decomposed into products such as CO2. Here we test underlying assumptions of the dominant conceptual model of litter decomposition. The model posits that a primary control on the rate of decomposition at regional to global scales is climate (temperature and moisture), with the controlling effects of decomposers negligible at such broad spatial scales. Using a regional-scale litter decomposition experiment at six sites spanning from northern Sweden to southern France—and capturing both within and among site variation in putative controls—we find that contrary to predictions from the hierarchical model, decomposer (microbial) biomass strongly regulates decomposition at regional scales. Furthermore, the size of the microbial biomass dictates the absolute change in decomposition rates with changing climate variables. Our findings suggest the need for revision of the hierarchical model, with decomposers acting as both local- and broad-scale controls on litter decomposition rates, necessitating their explicit consideration in global biogeochemical models.
  • Ecosphere

    Possible mechanisms underlying abundance and diversity responses of nematode communities to plant diversity

    Roeland Cortois, Ciska Veen, Henk Duyts, M. Abbas, Tanja Strecker, Olga Kostenko, Nico Eisenhauer, S. Scheu, G. Gleixner, Gerlinde De Deyn, Wim H. van der Putten
    Plant diversity is known to influence the abundance and diversity of belowground biota; however, patterns are not well predictable and there is still much unknown about the driving mechanisms. We analyzed changes in soil nematode community composition as affected by long-term manipulations of plant species and functional group diversity in a field experiment with plant species diversity controlled by sowing a range of 1–60 species mixtures and controlling non-sown species by hand weeding. Nematode communities contain a variety of species feeding on bacteria, fungi, plants, invertebrates, while some are omnivorous. We analyzed responses of nematode abundance and diversity to plant species and functional diversity, and used structural equation modeling (SEM) to explore the possible mechanisms underlying the observed patterns. The abundance of individuals of all nematode feeding types, except for predatory nematodes, increased with both plant species and plant functional group diversity. The abundance of microbial-feeding nematodes was related positively to aboveground plant community biomass, whereas abundance of plant-feeding nematodes was related positively to shoot C:N ratio. The abundance of predatory nematodes, in turn, was positively related to numbers of plant-feeding nematodes, but not to the abundance of microbial feeders. Interestingly, the numbers of plant-feeding nematodes per unit root mass were lowest in the high-diversity plant communities, pointing at reduced exposure to belowground herbivores when plants grow in species-diverse communities. Taxon richness of plant-feeding and microbial-feeding nematodes increased with plant species and plant functional group diversity. Increasing plant functional group diversity also enhanced taxon richness of predatory nematodes. The SEM suggests that bottom-up control effects of plant species and plant functional group diversity on abundance of nematodes in the various feeding types predominantly involve mechanistic linkages related to plant quality instead of plant quantity; especially, C:N ratios of the shoot tissues, and/or effects of plants on the soil habitat, rather than shoot quantity explained nematode abundance. Although aboveground plant properties may only partly serve as a proxy for belowground resource quality and quantity, our results encourage further studies on nematode responses to variations in plant species and plant functional diversity in relation to both quantity and quality of the belowground resources.
  • Oikos

    Coordinated responses of soil communities to elevation in three subarctic vegetation types

    Ciska Veen, Jonathan R. De Long, Paul Kardol, Maja K. Sundqvist, Basten Snoek, David A. Wardle
    Global warming has begun to have a major impact on the species composition and functioning of plant and soil communities. However, long-term community and ecosystem responses to increased temperature are still poorly understood. In this study, we used a well-established elevational gradient in northern Sweden to elucidate how plant, microbial and nematode communities shift with elevation and associated changes in temperature in three highly contrasting vegetation types (i.e. heath, meadow and Salix vegetation). We found that responses of both the abundance and composition of microbial and nematode communities to elevation differed greatly among the vegetation types. Within vegetation types, changes with elevation of plant, microbial and nematode communities were mostly linked at fine levels of taxonomic resolution, but this pattern disappeared when coarser functional group levels were considered. Further, nematode communities shifted towards more conservative nutrient cycling strategies with increasing elevation in heath and meadow vegetation. Conversely, in Salix vegetation microbial communities with conservative strategies were most pronounced at the mid-elevation. These results provide limited support for increasing conservative nutrient cycling strategies at higher elevation (i.e. with a harsher climate). Our findings indicate that climate-induced changes in plant community composition may greatly modify or counteract the impact of climate change on soil communities. Therefore, to better understand and predict ecosystem responses to climate change, it will be crucial to consider vegetation type and its specific interactions with soil communities.
  • Functional Ecology

    Where, when and how plant-soil feedback matters in a changing world

    Wim H. van der Putten, Mark Bradford, Pella Brinkman, Tess Van de Voorde, Ciska Veen
    It is increasingly acknowledged that plant-soil feedbacks may play an important role in driving the composition of plant communities and functioning of terrestrial ecosystems. However, the mechanistic understanding of plant-soil feedbacks, as well as their roles in natural ecosystems in proportion to other possible drivers, is still in its infancy. Such knowledge will enhance our capacity to determine the contribution of plant-soil feedback to community and ecosystem responses under global environmental change. Here, we review how plant-soil feedbacks may develop under extreme drought and precipitation events, CO2 and nitrogen enrichment, temperature increase, land use change and plant species loss vs. gain. We present a framework for opening the black box of soil' considering the responses of the various biotic components (enemies, symbionts and decomposers) of plant-soil feedback to the global environmental changes, and we discuss how to integrate these components to understand and predict the net effects of plant-soil feedbacks under the various scenarios of change. To gain an understanding of how plant-soil feedback plays out in realistic settings, we also use the framework to discuss its interaction with other drivers of plant community composition, including competition, facilitation, herbivory, and soil physical and chemical properties. We conclude that understanding the role that plant-soil feedback plays in shaping the responses of plant community composition and ecosystem processes to global environmental changes requires unravelling the individual contributions of enemies, symbionts and decomposers. These biotic factors may show different response rates and strengths, thereby resulting in different net magnitudes and directions of plant-soil feedbacks under various scenarios of global change. We also need tests of plant-soil feedback under more realistic conditions to determine its contribution to changes in patterns and processes in the field, both at ecologically and evolutionary relevant time-scales.
  • Aquatic Botany

    Herbivory on freshwater and marine macrophytes: a review and perspective

    (Liesbeth) E.S. Bakker, Kevin A. Wood, Jordi F. Pagès, Ciska Veen, Marjolijn J.A. Christianen, Luis Santamaría, Bart A. Nolet, Sabine Hilt
    Until the 1990s, herbivory on aquatic vascular plants was considered to be of minor importance, and the predominant view was that freshwater and marine macrophytes did not take part in the food web: their primary fate was the detritivorous pathway. In the last 25 years, a substantial body of evidence has developed that shows that herbivory is an important factor in the ecology of vascular macrophytes across freshwater and marine habitats. Herbivores remove on average 40-48% of plant biomass in freshwater and marine ecosystems, which is typically 5-10 times greater than reported for terrestrial ecosystems. This may be explained by the lower C:N stoichiometry found in submerged plants. Herbivores affect plant abundance and species composition by grazing and bioturbation and therewith alter the functioning of aquatic ecosystems, including biogeochemical cycling, carbon stocks and primary production, transport of nutrients and propagules across ecosystem boundaries, habitat for other organisms and the level of shoreline protection by macrophyte beds. With ongoing global environmental change, herbivore impacts are predicted to increase. There are pressing needs to improve our management of undesirable herbivore impacts on macrophytes (e.g. leading to an ecosystem collapse), and the conflicts between people associated with the impacts of charismatic mega-herbivores. While simultaneously, the long-term future of maintaining both viable herbivore populations and plant beds should be addressed, as both belong in complete ecosystems and have co-evolved in these long before the increasing influence of man. Better integration of the freshwater, marine, and terrestrial herbivory literatures would greatly benefit future research efforts.
  • New Phytologist

    Effects of root decomposition on plant–soil feedback of early– and mid–successional plant species.

    Plant–soil feedback (PSF) is an important driver of plant community dynamics. Many studies have emphasized the role of pathogens and symbiotic mutualists in PSFs; however, less is known about the contribution of decomposing litter, especially that of roots.
    We conducted a PSF experiment, where soils were conditioned by living early- and mid-successional grasses and forbs with and without decomposing roots of conspecific species (conditioning phase). These soils were used to test growth responses of conspecific and heterospecific plant species (feedback phase).
    The addition of the roots of conspecifics decreased the biomass of both early- and mid-successional plant species in the conditioning phase. In the feedback phase, root addition had positive effects on the biomass of early-successional species and neutral effects on mid-successional species, except when mid-successional grasses were grown in soils conditioned by conspecifics, where effects were negative. Biomass of early- and mid-successional forbs was generally reduced in soils conditioned by conspecifics.
    We conclude that root decomposition may increase short-term negative PSF effects, but that the effects can become neutral to positive over time, thereby counteracting negative components of PSF. This implies that root decomposition is a key element of PSF and needs to be included in future studies.
  • Oikos

    Litter quality and environmental controls of home-field advantage effects on litter decomposition

    Ciska Veen, G.T. Freschet, A. Ordonez, David A. Wardle
    The ‘home-field advantage (HFA) hypothesis’ predicts that plant litter is decomposed faster than expected in the vicinity of the plant where it originates from (i.e. its ‘home’) relative to some other location (i.e. ‘away’) because of the presence of specialized decomposers. Despite growing evidence for the widespread occurrence HFA effects, what drives HFA is not understood as its strength appears highly variable and context-dependent. Our work advances current knowledge about HFA effects by testing under what conditions HFA is most important. Using published data on mass loss from 125 reciprocal litter transplants from 35 studies, we evaluated if HFA effects were modulated by macroclimate, litter quality traits, and the dissimilarity between ‘home’ and ‘away’ of both the quality of reciprocally exchanged litters and plant community type. Our results confirmed the occurrence of an overall, worldwide, HFA effect on decomposition with on average 7.5% faster decomposition at home. However, there was considerable variation in the strength and direction (sometimes opposite to expectations) of these effects. While macroclimate and average litter quality had weak or no impact on HFA effects, home-field effects became stronger (regardless of the direction) when the quality of ‘home’ and ‘away’ litters became more dissimilar (e.g. had a greater dissimilarity in N:P ratio; F1,42 = 6.39, p = 0.015). Further, home-field effects were determined by the degree of difference between the types of dominant plant species in the ‘home’ versus ‘away’ communities (F2,105 = 4.03, p = 0.021). We conclude that home-field advantage is not restricted to particular litter types or climate zones, and that the dissimilarity in plant communities and litter quality between the ‘home’ and ‘away’ locations, are the most significant drivers of home-field effects.
  • Oikos

    Plant growth response to direct and indirect temperature effects varies by vegetation type and elevation in a subarctic tundra

    Jonathan R. De Long, Paul Kardol, Maja K. Sundqvist, Ciska Veen, David A. Wardle
    There has been growing recent use of elevational gradients as tools for assessing effects of temperature changes on vegetation properties, because these gradients enable temperature effects to be considered over larger spatial and temporal scales than is possible through conventional experiments. While many studies have explored the direct effects of temperature, the indirect effects of temperature through its long-term influence on soil abiotic or biotic properties remain essentially unexplored. We performed two climate chamber experiments using soils from a subarctic elevational gradient in Abisko, Sweden to investigate the direct effects of temperature, and indirect effects of temperature via soil legacies, on growth of two grass species. The soils were collected from each of two vegetation types (heath, dominated by dwarf shrubs, and meadow, dominated by graminoids and herbs) at each of three elevations. We found that plants responded to both the direct effect of temperature and its indirect effect via soil legacies, and that direct and indirect effects were largely decoupled. Vegetation type was a major determinant of plant responses to both the direct and indirect effects of temperature; responses to soils from increasing elevation were stronger and showed a more linear decline for meadow than for heath soils. The influence of soil biota on plant growth was independent of elevation, with a positive influence across all elevations regardless of soil origin for meadow soils but not for heath soils. Taken together, this means that responses of plant growth to soil legacy effects of temperature across the elevational gradient were driven primarily by soil abiotic, and not biotic, factors. These findings emphasize that vegetation type is a strong determinant of how temperature variation across elevational gradients impacts on plant growth, and highlight the need for considering both direct and indirect effects of temperature on plant responses to future climate change.
  • New Phytologist

    Peeking into the black box: A trait-based approach to predicting plant-soil feedback

    Paul Kardol, Ciska Veen, François P. Teste, M.P. Perring
    Feedbacks between plants and soil communities may be elusive, yet
    they have far-reaching consequences for plant physiology, competition
    and community structure. Plant–soil feedbacks (PSFs) are
    plant-mediated changes to soil properties that ultimately influence
    the performance of the same or other plants (Van der Putten et al.,
    2013). These PSFs may be mediated by root-associated organisms
    (hereafter, root-mediated feedbacks) or saprotrophic organisms
    and associated litter characteristics (hereafter, litter-mediated
    feedbacks). However, we know little about the potential mechanistic
    linkages and relative strengths between these distinct, but
    connected, processes as root- and litter-mediated feedbacks have
    generally been studied independently from each other. This is
    despite the fact that root-associated organisms and saprotrophs can
    interact through various mechanisms, either directly or as mediated
    by the plant (e.g. Wardle, 2006). By using a trait-based approach,Ke
    et al. (in this issue of New Phytologist, pp. 329–341) make an
    important contribution by integrating root- and litter-mediated
    PSFs in a nitrogen (N)-based, stage-structured plant population
    and microbial community model. Their approach allows us to start
    peeking into the ‘black box’ thereby promoting a better understanding
    of how PSFs operate interactively. Ke et al. considered
    various plant traits (e.g. decomposability), but also incorporated
    trait variation in the physiology, demography and composition of
    the soil microbial community, and tested their separate and
    interactive effects on PSF strength in a comprehensive simulation
    framework. Finally, they used empirical evidence from the
    literature to support their model predictions.
  • Functional Ecology

    Environmental factors and traits that drive plant litter decomposition do not determine home-field advantage effects

    Ciska Veen, Maja K. Sundqvist, David A. Wardle
    The ‘home-field advantage’ (HFA) hypothesis predicts that plant litter is decomposed faster than expected underneath the plant from which it originates (‘home’) than underneath other plants (‘away’), because decomposer communities are specialized to break down litter from the plants they associate with. However, empirical evidence shows that the occurrence of HFA is highly variable, and the reasons for this are little understood. In our study we progress our understanding by investigating whether HFA is stronger for more recalcitrant litter types and under colder conditions and how soil properties and plant functional traits affect the magnitude and direction of HFA. In subarctic tundra in northern Sweden we set up a reciprocal transplant litter decomposition experiment along an elevational gradient where three highly contrasting vegetation types (heath, meadow and Salix) occur at all elevations, and where temperature decreases strongly with elevation. In this study, we used a litter bag approach where litters from each elevation × vegetation type combination were decomposed in all combinations of elevation × vegetation type. We also measured community-level plant functional traits, such as leaf and litter nutrient content. We determined soil biotic and abiotic properties, such as microbial biomass and soil nutrient content, in soil cores collected for each elevation × vegetation type combination. We found that mass loss increased with plant and litter nutrient content and with soil temperature. In contrast, the occurrence of HFA was limited in our study system, and its magnitude and direction could not be explained by vegetation type, elevation, plant traits or soil properties, despite these factors serving as powerful drivers of litter mass loss in our study. We conclude that although vegetation type and climate are major drivers of litter mass loss, they do not emerge as important determinants of HFA. Therefore, while rapid shifts in plant community composition or temperature due to global change are likely to influence litter mass loss directly by altering environmental conditions, plant trait spectra and litter quality, indirect effects of global change resulting from decoupling of specialist interactions between litter and decomposer communities appears to be of less importance. This article is protected by copyright. All rights reserved.
  • Arctic, Antarctic, and Alpine Research

    Above-ground and below-ground plant responses to fertilization in two subarctic ecosystems

    Ciska Veen, Maja K. Sundqvist, D. Metcalfe, S.D. Wilson
    Soil nutrient supply is likely to change in the Arctic due to altered process rates associated with climate change. Here, we compare the responses of herbaceous tundra and birch forest understory to fertilization, considering both above- and below-ground responses. We added nitrogen and phosphorus to plots in both vegetation types for three years near Abisko, northern Sweden, and measured the effect on above- and below-ground plant community properties and soil characteristics. Fertilization increased ground-layer shoot mass, the cover of grasses, and tended to enhance total root length below-ground, while it reduced the cover of low statured deciduous dwarf-shrubs. The only statistically significant interaction between vegetation type and fertilization was for grass cover, which increased twofold in forest understory but sixfold in tundra following fertilization. The lack of interactions for other variables suggests that the ground layers in these contrasting vegetation types have similar responses to fertilization. The nutrient-driven increase in grass cover and species-specific differences in productivity and root characters may alter ecosystem dynamics and C cycling in the long-term, but our study indicates that the response of birch forest understory and tundra vegetation may be consistent.
  • Ecosystems

    Herbivores enforce sharp boundaries between terrestrial and aquatic ecosystems

    Judith Sarneel, Naomi Huig, Ciska Veen, W. Rip, (Liesbeth) E.S. Bakker
    The transitions between ecosystems (ecotones) are often biodiversity hotspots, but we know little about the forces that shape them. Today, often sharp boundaries with low diversity are found between terrestrial and aquatic ecosystems. This has been attributed to environmental factors that hamper succession. However, ecosystem properties are often controlled by both bottom-up and top-down forces, but their relative importance in shaping riparian boundaries is not known. We hypothesize that (1) herbivores may enforce sharp transitions between terrestrial and aquatic ecosystems by inhibiting emergent vegetation expansion and reducing the width of the transition zone and (2) the vegetation expansion, diversity, and species turnover are related to abiotic factors in the absence of herbivores, but not in their presence. We tested these hypotheses in 50 paired grazed and ungrazed plots spread over ten wetlands, during two years. Excluding grazers increased vegetation expansion, cover, biomass, and species richness. In ungrazed plots, vegetation cover was negatively related to water depth, whereas plant species richness was negatively related to the vegetation N:P ratio. The presence of (mainly aquatic) herbivores overruled the effect of water depth on vegetation cover increase but did not interact with vegetation N:P ratio. Increased local extinction in the presence of herbivores explained the negative effect of herbivores on species richness, as local colonization rates were unaffected by grazing. We conclude that (aquatic) herbivores can strongly inhibit expansion of the riparian vegetation and reduce vegetation diversity over a range of environmental conditions. Consequently, herbivores enforce sharp boundaries between terrestrial and aquatic ecosystems.
  • Oikos

    Grazing-induced changes in plant–soil feedback alter plant biomass allocation

    Large vertebrate herbivores, as well as plant–soil feedback interactions are important drivers of plant performance, plant community composition and vegetation dynamics in terrestrial ecosystems. However, it is poorly understood whether and how large vertebrate herbivores and plant–soil feedback effects interact. Here, we study the response of grassland plant species to grazing-induced legacy effects in the soil and we explore whether these plant responses can help us to understand long-term vegetation dynamics in the field. In a greenhouse experiment we tested the response of four grassland plant species, Agrostis capillaris, Festuca rubra, Holcus lanatus and Rumex acetosa, to field-conditioned soils from grazed and ungrazed grassland. We relate these responses to long-term vegetation data from a grassland exclosure experiment in the field. In the greenhouse experiment, we found that total biomass production and biomass allocation to roots was higher in soils from grazed than from ungrazed plots. There were only few relationships between plant production in the greenhouse and the abundance of conspecifics in the field. Spatiotemporal patterns in plant community composition were more stable in grazed than ungrazed grassland plots, but were not related to plant–soil feedbacks effects and biomass allocation patterns. We conclude that grazing-induced soil legacy effects mainly influenced plant biomass allocation patterns, but could not explain altered vegetation dynamics in grazed grasslands. Consequently, the direct effects of grazing on plant community composition (e.g. through modifying light competition or differences in grazing tolerance) appear to overrule indirect effects through changes in plant–soil feedback
  • Theoretical Ecology

    Plant-soil feedbacks and the coexistence of competing plants

    T.A. Revilla, Ciska Veen, M.B. Eppinga, F.J. Weissing
    Plant–soil feedbacks can have important implications for the interactions among plants. Understanding these effects is a major challenge since it is inherently difficult to measure and manipulate highly diverse soil communities. Mathematical models may advance this understanding by making the interplay of the various processes affecting plant–soil interaction explicit and by quantifying the relative importance of the factors involved. The aim of this paper is to provide a complete analysis of a pioneering plant–soil feedback model developed by Bever and colleagues (J Ecol 85: 561–573, 1997; Ecol Lett 2: 52–62, 1999; New Phytol 157: 465–473, 2003) to fully understand the range of possible impacts of plant–soil feedbacks on plant communities within this framework. We analyze this model by means of a new graphical method that provides a complete classification of the potential effects of soil communities on plant competition. Due to the graphical character of the method, the results are relatively easy to obtain and understand. We show that plant diversity depends crucially on two key parameters that may be viewed as measures of the intensity of plant competition and the direction and strength of plant–soil feedback, respectively. Our analysis provides a formal underpinning of earlier claims that plant–soil feedbacks, especially when they are negative, may enhance the diversity of plant communities. In particular, negative plant–soil feedbacks can enhance the range of plant coexistence by inducing competitive oscillations. However, these oscillations can also destabilize plant coexistence, leading to low population densities and extinctions. In addition, positive feedbacks can allow locally stable forms of plant coexistence by inducing alternative stable states. Our findings highlight that the inclusion of plant–soil interactions may fundamentally alter the predictions on the structure and functioning of above-ground ecosystems. The scenarios presented in this study can be used to formulate hypotheses about the ways soil community effects may influence plant competition that can be tested with empirical studies. This will advance our understanding of the role of plant–soil feedback in ecological communities.
  • Freshwater Biology

    Aquatic grazers reduce the establishment and growth of riparian plants along an environmental gradient

    Ciska Veen, Judith Sarneel, Lone Ravensbergen, Naomi Huig, José van Paassen, W. Rip, (Liesbeth) E.S. Bakker
    Summary The establishment of riparian plants is determined by abiotic conditions and grazing, although it is usually presumed that the former are most important. We tested the impact of aquatic grazers on the survival and growth of establishing riparian plants and whether the impact of grazing interacts with abiotic conditions. We conducted an experiment across 10 Dutch wetlands, covering a large range of water depth and nutrient availability. We introduced 1-year-old plants of an emergent (common reed, Phragmites australis) and a floating (water soldier, Stratiotes aloides) species in individual enclosures (n = 5 per site) that excluded predominantly waterbirds, which were the most abundant grazers, and on adjacent unprotected plots. Survival and growth were measured during one growing season. Grazing reduced growth (as biomass) of Phragmites and Stratiotes by a mean of 25 and 60%, respectively. Grazing decreased survival of Stratiotes, but not of Phragmites. Shallow water, water-level fluctuations, eutrophic conditions and enough light favoured both growth and survival of Phragmites. Growth of Stratiotes was unaffected by these factors, but they reduced its survival. For both species, grazing effects on biomass were consistent across environmental conditions, but for Phragmites, grazing effects on survival were influenced by abiotic conditions. We conclude that aquatic grazers significantly reduce the establishment and growth of macrophytes in the riparian zone over a wide range of environmental conditions.
  • Perspectives in Plant Ecology, Evolution and Systematics

    An integrated perspective to explain nitrogen mineralization in grazed ecosystems

    Maarten Schrama, Ciska Veen, (Liesbeth) E.S. Bakker, J.L. Ruifrok, Jan P. Bakker, H. Olff
    Large herbivores are key drivers of nutrient cycling in ecosystems worldwide, and hence they have an important influence on the productivity and species composition in plant communities. Classical theories describe that large herbivores can accelerate or decelerate nitrogen (N) mineralization by altering the quality and quantity of resource input (e.g. dung, urine, plant litter) into the soil food web. However, in many situations the impact of herbivores on N mineralization cannot be explained by changes in resource quality and quantity. In this paper, we aim to reconcile observations of herbivores on N mineralization that were previously regarded as contradictory. We conceptually integrate alternative pathways via which herbivores can alter N mineralization. We illustrate our new integrated perspective by using herbivore-induced soil compaction and subsequent changes in soil moisture and soil aeration as an example. We show that the net effect of herbivores on mineralization depends on the balance between herbivore-induced changes in soil physical properties and changes in the quality and quantity of resource input into the soil food web. For example, soil compaction by herbivores can limit oxygen or water availability in wet and dry soils respectively, particularly those with a fine texture. This can result in a reduction in N mineralization regardless of changes in resource quality or quantity. In such systems the plant community will shift towards species that are adapted to waterlogging (anoxia) or drought, respectively. In contrast, soils with intermediate moisture levels are less sensitive to compaction. In these soils, N mineralization rates are primarily associated with changes in resource quality and quantity. We conclude that our integrated perspective will help us to better understand when herbivores accelerate or decelerate soil nutrient cycling and improve our understanding of the functioning of grazed ecosystems
  • Oecologia

    Large grazers modify effects of aboveground–belowground interactions on small-scale plant community composition

    Ciska Veen, E. Geuverink, H. Olff
    Aboveground and belowground organisms influence plant community composition by local interactions, and their scale of impact may vary from millimeters belowground to kilometers aboveground. However, it still poorly understood how large grazers that select their forage on large spatial scales interact with small-scale aboveground– belowground interactions on plant community heterogeneity. Here, we investigate how cattle (Bos taurus) modify the effects of interactions between yellow meadow ants (Lasius flavus) and European brown hares (Lepus europaeus) on the formation of small-scale heterogeneity in vegetation composition. In the absence of cattle, hares selectively foraged on ant mounds, while under combined grazing by hares and cattle, vertebrate grazing pressure was similar on and off mounds. Ant mounds that were grazed by only hares had a different plant community composition compared to their surroundings: the cover of the grazingintolerant grass Elytrigia atherica was reduced on ant mounds, whereas the relative cover of the more grazingtolerant and palatable grass Festuca rubra was enhanced. Combined grazing by hares and cattle, resulted in homogenization of plant community composition on and off ant mounds, with high overall cover of F. rubra. We conclude that hares can respond to local ant–soil–vegetation interactions, because they are small, selective herbivores that make their foraging decisions on a local scale. This results in small-scale plant patches on mounds of yellow meadow ants. In the presence of cattle, which are less selective aboveground herbivores, local plant community patterns triggered by small-scale aboveground– belowground interactions can disappear. Therefore, cattle modify the consequences of aboveground–belowground interactions for small-scale plant community composition.
  • Basic and Applied Ecology

    Interactive effects of soil-dwelling ants, ant mounds and simulated grazing on local plant community composition

    Ciska Veen, H. Olff
    Interactions between aboveground vertebrate herbivores and subterranean yellow meadow ants (Lasius flavus) can drive plant community patterns in grassland ecosystems. Here, we study the relative importance of the presence of ants (L. flavus) and ant mounds under different simulated grazing regimes for biomass production and species composition in plant communities. We set up a greenhouse experiment using intact soil cores with their associated vegetation. We found that plant biomass production in the short term was affected by an interaction between simulated grazing (clipping) and ant mound presence. Clipping homogenized production on and off mounds, while in unclipped situations production was higher off than on mounds. During the experiment, these differences in unclipped situations disappeared, because production on unclipped mounds increased. Plant species richness was on average higher in clipped treatments and patterns did not change significantly over the experimental period. Plant community composition was mainly affected by clipping, which increased the cover of grazing-tolerant plant species. The actual presence of yellow meadow ants did not affect plant community composition and production. We conclude that the interaction between ant mounds and clipping determined plant community composition and biomass production, while the actual presence of ants themselves was not important. Moreover, clipping can overrule effects of ant mounds on biomass production. Only shortly after the cessation of clipping biomass production was affected by ant mound presence, suggesting that only under low intensity clipping ant mounds may become important determining plant production. Therefore, under low intensity grazing ant mounds may drive the formation of small-scale plant patches.
  • Ecology

    Vertebrate herbivores influence soil nematodes by modifying plant communities

    Ciska Veen, H. Olff, Henk Duyts, Wim H. van der Putten
    Abiotic soil properties, plant community composition, and herbivory all have been reported as important factors influencing the composition of soil communities. However, most studies thus far have considered these factors in isolation, whereas they strongly interact in the field. Here, we study how grazing by vertebrate herbivores influences the soil nematode community composition of a floodplain grassland while we account for effects of grazing on plant community composition and abiotic soil properties. Nematodes are the most ubiquitous invertebrates in the soil. They include a variety of feeding types, ranging from microbial feeders to herbivores and carnivores, and they perform key functions in soil food webs. Our hypothesis was that grazing affects nematode community structure and composition through altering plant community structure and composition. Alternatively, we tested whether the effects of grazing may, directly or indirectly, run via changes in soil abiotic properties. We used a long-term field experiment containing plots with and without vertebrate grazers (cattle and rabbits). We compared plant and nematode community structure and composition, as well as a number of key soil abiotic properties, and we applied structural equation modeling to investigate four possible pathways by which grazing may change nematode community composition. Aboveground grazing increased plant species richness and reduced both plant and nematode community heterogeneity. There was a positive relationship between plant and nematode diversity indices. Grazing decreased the number of bacterial-feeding nematodes, indicating that in these grasslands, top-down control of plant production by grazing leads to bottom-up control in the basal part of the bacterial channel of the soil food web. According to the structural equation model, grazing had a strong effect on soil abiotic properties and plant community composition, whereas plant community composition was the main determinant of nematode community composition. Other pathways, which assumed that grazing influenced nematode community composition by inducing changes in soil abiotic properties, did not significantly explain variation in nematode community composition. We conclude that grazing-induced changes in nematode community composition mainly operated via changes in plant community composition. Influences of vertebrate grazers on soil nematodes through modification of abiotic soil properties were of less importance.
  • Oikos

    Influence of grazing and fire frequency on small-scale plant community structure and resource variability in native tallgrass prairie

    Ciska Veen, John Blair, Melinda D. Smith, Scott L. Collins
    In grasslands worldwide, grazing by ungulates and periodic fires are important forces affecting resource availability and plant community structure. It is not clear, however, whether changes in community structure are the direct effects of the disturbance (i.e. fire and grazing) or are mediated indirectly through changes in resource abundance and availability. In North American tallgrass prairies, fire and grazing often have disparate effects on plant resources and plant diversity, yet, little is known about the individual and interactive effects of fire and grazing on resource variability and how that variability relates to heterogeneity in plant community structure, particularly at small scales. We conducted a field study to determine the interactive effects of different long-term fire regimes (annual vs four-year fire frequency) and grazing by native ungulates (Bos bison) on small-scale plant community structure and resource variability (N and light) in native tallgrass prairie. Grazing enhanced light and nitrogen availability, but did not affect small-scale resource variability. In addition, grazing reduced the dominance of C4 grasses which enhanced species richness, diversity and community heterogeneity. In contrast, annual fire increased community dominance and reduced species richness and diversity, particularly in the absence of grazing, but had no effect on community heterogeneity, resource availability and resource variability. Variability in the abundance of resources showed no relationship with community heterogeneity at the scale measured in this study, however we found a relationship between community dominance and heterogeneity. Therefore, we conclude that grazing generated small-scale community heterogeneity in this mesic grassland by directly affecting plant community dominance, rather than indirectly through changes in resource variability.

Projecten & samenwerkingen


  • Carbon storage

    Project 2022
    Carbon storage
  • Nutrient Network (NutNet)

    Project 2017–Present
    Vertebrate herbivores and nutrient deposition have a strong impact on the biodiversity and functioning of grasslands worldwide. Both factors have been changing rapidly over the last decades due to exctinction of herbivores, restoration of herbivore communities (e.g., rewilding) and enhanced nutrient inputs in natural grasslands via agriculture and fossil fuel combustion. In a global network of experiments we quantify how these changse impact on plant, soil and insect biodiversity, the cycling of nutrients and other functions in grassland ecosystems ( At the NIOO, we coordinate the Dutch NutNet site, which is situated on the Veluwe.
    NutNet site
  • Climate-Smart Forests

    Project 2021–2026
    As a response to global climate change, which is putting increased pressure on most ecosystems, national and international agreements aim at creating forests that are productive, resilient to climate change, and that store carbon to mitigate global warming. However, these aims are being challenged by increased tree mortality rates and decreased tree growth rates in response to increased incidence of drought. The summer drought of 2018 alone resulted in 100 million m3 of dead trees in Europe, equivalent to a loss of approximately 3.5 billion euros wood.
    Therefore, the challenge is to develop climate-smart forestry (CSF) in order to sustain or increase forest productivity, forest resilience and forest carbon storage under climate change. Currently, there is a lack of crucial insights into the effects of forest management on the growth and survival of trees, and on carbon storage in both trees and forest soils, particularly under increased incidence of drought. We test the hypothesis that CSF aims can be achieved via controlling stand density by applying intermediate levels of tree harvest intensity. The main aim of this proposed project is to quantify the effects of drought and management-controlled stand density on forest productivity, forest resilience, and carbon storage in trees and soils.
    Beech forests in the Netherlands
  • Soil biodiversity and functioning in food forests

    Project 2020–2025
    Temperate food forests have gained attention over the last decade because of their potential to contribute to restoration of biodiversity and carbon storage. So far scientific research has been limited to case studies and identifying socio-economic values. In this project, we aim to understand how food forestry affects belowground biodiversity and functioning (including carbon storage) compared to other types of land use (e.g., arable farming, grassland). We collect field observations and use controlled experiments. This project is part of a larger TKI program, where also aboveground biodiversity and earning models of food forests are investigated. For information see:
    Field work in the food forest
  • Vital soils for sustainable intensification of agriculture

    Project 2016–2021
    A key challenge for sustainable intensification of agriculture is to produce increasing amounts of food for a growing world population, with minimal loss of biodiversity and ecosystem services. In order to facilitate ecological intensification of agriculture, the underlying principles need to be understood and validated in farmers’ fields

Additionele Projecten

  • NWO-Veni


    Testing the role of specialized microbial communities in driving home-field advantage effects (i.e., accelerated litter breakdown near plants where litter originates from) for litter decomposition.

  • NWO-Rubicon


    Testing home-field advantage (i.e., accelerated litter breakdown near plants where litter originates from) for litter decomposition along climate and litter quality gradients.