, Volume 189, Issue 3, pp 781–793 | Cite as

Responses of seed size, ant worker size, and seed removal rate to elevation in Mediterranean grasslands

  • Mariola SilvestreEmail author
  • Arantxa Aguilar
  • Javier Seoane
  • Francisco M. Azcárate
Community ecology – original research


Elevational gradients are a powerful tool to investigate how abiotic factors affect ecosystems and to predict the possible consequences of climate change. Here, we investigated the assemblage-level responses of seed size, ant worker size, and rates of seed removal by ants along a 1700 m elevational gradient in Mediterranean grasslands of central Spain, taking into account abiotic factors and interactions between biotic variables. Our results showed that both seed size and ant size decreased with elevation. Structural equation models linking abiotic factors and biotic variables were built to describe these patterns. Two equally plausible models were selected, both based on temperature. In the first model, temperature directly affected seed size and granivorous ant size, and indirectly affected ant size (of both granivorous and non-granivorous species) through a bottom-up pathway. The second model included the direct effects of temperature on ant size and granivorous ant size, and indirect effects on seed size through a top-down pathway. In this model, the link between granivorous ant size and seed size was positive. Seed removal rates decreased with elevation and seed size, with smaller seeds being preferred with increasing elevation. Taken together, our results indicate that the observed patterns of ant size and seed size are ultimately controlled by abiotic factors, although the system is more satisfactorily explained if seed–ant interactions are incorporated into the models. We expect that climate change will have strong effects on this system, including mismatches between consumers and producers potentially altering the interspecific relationships among ants and plants.


Altitude Climate change Granivory Harvester ants Seed predation 



Carried out within a chronically low-budgeted science system struck by an economic crisis, this study could not have been made without the enthusiastic support of students and friends, who helped with field work (Sonia Hernando, Rubén Ariño, Julia Gómez, Adrián Barrero, A.E. Santamaría, Adrián Lorenzo, Montse Reina, Esperanza Iranzo, Noelia González, Miguel Muñoz, Violeta Hevia, Cristina Rota, Manuel Rojo, and Javier Cañas) and satellite data analyses (Miguel Blázquez). Xavier Picó kindly reviewed a preliminary version of the paper. However, we benefited from the administrative and financial support by the Spanish MINECO (CGL2014-53789-R) and Sierra de Guadarrama National Park. MS, JS, and FMA work within the Madrid’s Government research group network REMEDINAL3-CM (S-2013/MAE-2719). MS was supported by an FPI grant from MINECO. Nuria Plaza, from the Instituto Geográfico Nacional (, kindly provided access to and help with the satellite imagery.

Author contribution statement

JS and FMA originally formulated the idea. MS, AA, JS, and FMA designed the study, conducted field work, and analyzed the data. MS, JS, and FMA wrote the manuscript.

Supplementary material

442_2019_4356_MOESM1_ESM.pdf (212 kb)
Supplementary material 1 (PDF 211 kb)
442_2019_4356_MOESM2_ESM.pdf (171 kb)
Supplementary material 2 (PDF 171 kb)
442_2019_4356_MOESM3_ESM.pdf (157 kb)
Supplementary material 3 (PDF 157 kb)
442_2019_4356_MOESM4_ESM.pdf (394 kb)
Supplementary material 4 (PDF 393 kb)


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.Terrestrial Ecology Group, Department of EcologyUniversidad Autónoma de MadridMadridSpain
  2. 2.Centro de Investigación en Biodiversidad y Cambio Global (CIBC-UAM), Universidad Autónoma de MadridMadridSpain

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