Dr. Rainer Wirth

Senior Scientist & Lecturer


Room: 70/219
Phone: 0631 - 205 4401
Mail: wirth(at)rhrk.uni-kl.de


Molecular Botany (Plant Ecology Lab)
Faculty of Biology
University of Kaiserslautern-Landau
PO-BOX 3049
7653 Kaiserslautern


Research interests

Species interactions under human pressure

Today's landscapes are subjected to unprecedented rates of human-induced habitat modifications resulting in a severe ecosystem transformation including the re-assemblage of biotic communities, the alteration of vegetation dynamics, habitat simplification, and the alteration of species interactions. Our interest is aimed at (i) understanding the contemporary forces driving the facilitation or distortion of species interactions, (ii) describing the nature of these changes and (iii) exploring the multifaceted feedbacks on biodiversity, ecosystem stability and functioning in both temperate and tropical forested systems. Ongoing research activities include:

  • Impacts of silvicultural land use and forest fragmentation on herbivore performance and plant-herbivore interactions (networks) in a temperate forest landscape
  • Proliferation of disturbance-adapted organisms and biodiversity prospects in fragmented forest landscapes
  • Trophic interactions in tropical forest fragments

See >> here for further information on current projects


Interactions between plants and herbivores are considered key processes shaping life history traits and ecosystem function. By consuming plant tissues, herbivores are able to alter plant performance, reduce plant fitness, and influence ecosystem functions. On the other hand evolved plant traits, phenotypic variation, and inducible responses strongly influence the performance, behavioural patterns, and community composition of herbivores. Our main research efforts are dedicated to:

  • Damage and foraging patterns of herbivorous insects
  • Herbivory impacts on plant performance (from population to ecosystem level)
  • Chemical ecology of food plant selection

Leaf cutting ant ecology

Leaf-cutting ants are very peculiar due to their unique ability to cultivate fungus for food and the many ecological implications resulting from this symbiosis. At the same time they are amongst the most polyphagous and voracious herbivorous insects worldwide cutting up to 15% of the standing leaf crop in their colony’s territory each year (Wirth et al. 2003, Urbas et al. 2007) and serve as key model organisms for investigating generalist herbivores. In addition we have accumulated evidence, that they substantially impact the functioning and the successional trajectory of disturbed neotropical forests via trophic activities (herbivory and seed dispersion, Silva et al. 2007, Bieber et al. 2011) and ecosystem engineering (Corrêa et al. 2010, Meyer et al. 2011a, 2011b). We work on several facets of leaf-cutting ant ecology including:

  • How leaf-cutting ants deal with and are affected by plant-induced defensive volatiles
  • Complexity and spatio-temporal variability of foraging trail networks in leaf-cutting ants
  • Leaf-cutting ants as winner species (sensu McKinney & Lockwood 1999) in human-modified ecosystems
  • Ecosystem engineering via nest-mediated shifts in forest microclimate and edaphic conditions
  • Consumption and herbivory levels by Atta spp.
  • Microbial and chemical ecology in the leaf-cutting ant cosmos (i.e. ants, fungus garden, and microbial antagonists/mutualists)

Academic services

  • Scientific Board/Subject Editor Biotropica, 2007- 2010
  • Scientific Board Society of Tropical Ecology (GTÖ), 2009-2014
  • Referee for: Biotropica, Oecologia, Ecology, Journal of Applied Entomology, Journal of Applied  Ecology, Behavioral Ecology, Studies on Neotropical Fauna and Environment, Journal of Ecology, Oikos, Journal of Chemical Ecology
  • Grant Reviewer for: Deutsche Forschungsgemeinschaft (DFG), Alexander von Humboldt Foundation, Czech Science Foundation, The International Foundation for Science, Rufford Foundation, The International Max Planck Research School (IMPRS), Fulbright Student Program.


  • Plant Ecology
  • Species Interactions
  • Biodiversity
  • Native Plant Identification
  • The Scientific Process


For current courses offered by the department of Molecular Botany see  >> here


Oliveira, F.R., Centeno-Alvarado, D., Oliveira, F.M.P., Wirth, R., Lopes, A.V., Leal, I.R. 2023 Divergent effects of leaf-cutting ant herbivory and soil engineering on the reproductive success of plants in a Caatinga dry forest. Ecological Entomology, DOI: 10.1111/een.13289

Szyja, M., Felde, V. J. M. N. L., Lückel, S., Tabarelli, M., Leal, I. R., Büdel, B. and Wirth, R. 2023. Biological soil crusts decrease infiltration but increase erosion resistance in a human-disturbed tropical dry forest. Frontiers in Microbiology 14, DOI:10.3389/fmicb.2023.1136322

Oliveira, F.M.P., Knoechelmann, C.M, Wirth, R., Tabarelli, M., Leal, I.R. 2023. Leaf-cutting ant nests support less dense and impoverished seed assemblages in a human-modified Caatinga dry forest. Biotropica, DOI: 10.1111/btp.13198

Noutcheu R., Oliveira F. M. P., Wirth R., Tabarelli M. and Leal I. R. 2023. Coppicing as a driver of plant resprouting and the regeneration of a Caatinga dry forest. Forest Ecology and Management,  DOI: 10.1016/j.foreco.2022.120736

Lins, L., Santos, D., Lira, R., Oliveira, F. M., Wirth, R., Menezes, T., Tabarelli, M.Leal, I. R. 2022. Exotic goats do not affect the seed bank but reduce seedling survival in a human-modified landscape of Caatinga dry forest. Forest Ecology and Management, 522, 120491. DOI: 10.1016/j.foreco.2022.120491

Santos-Neto P. E., Arnan X., Ribeiro-Neto J. D., Wirth R. and Leal I. R. 2022. Aridity, but not disturbance, reduces the specialization and modularity of plant-insect herbivore interaction networks in Caatinga dry forest. Journal of Insect Conservation, DOI: 10.1007/s10841-022-00376-5.

Barros, M., Ribeiro, E. M., Vanderlei, R., de Paula, A., Silva, A., Wirth, R., Cianciaruso, M. and Tabarelli, M. 2020. Resprouting drives successional pathways and the resilience of Caatinga dry forest in human-modified landscapes. Forest Ecology and Management, doi.org/10.1016/j.foreco.2020.118881

Menezes T., Carmo R., Wirth R., Leal I.R., Tabarelli M., Laurenio A., Melo F.P.L. 2020. Introduced goats reduce diversity and biomass of herbs in Caatinga dry forest. Land Degradation & Development, DOI: 10.1002/ldr.3693

Baehner K.W., Tabarelli M., Buedel B., Wirth R. 2020. Habitat fragmentation and forest management alter woody plant communities in a Central European beech forest landscape. Biodiversity and Conservation, DOI:10.1007/s10531-020-01996-6

Bustamante S., Amarillo-Suarez A.R., Wirth R. 2020. Effects of pasture and forest microclimatic conditions on the foraging activity of leaf-cutting ants. Biotropica, DOI:10.1111/btp.12783

Knoechelmann, C.M., Oliveira, F.M.P., Siqueira, F.F.S., Wirth, R., Tabarelli, M., Leal, I.R. 2020. Leaf-cutting ants negatively impact the regeneration of the Caatinga dry forest across abandoned pastures. Biotropica, DOI: 10.1111/btp.12782

Szyja, M., Menezes, A.G.S., Oliveira, F.D.A., Leal, I., Tabarelli, M., Buedel, B., Wirth, R. 2019. Neglected but Potent Dry Forest Players: Ecological role and ecosystem service provision of biological soil crusts in the human-modified Caatinga. Frontiers in Ecology and Evolution DOI: 10.3389/fevo.2019.00482

Ribeiro, E.M.S., Specht, M.J., Tabarelli, M., Santos, B.A., Gerhold, P., Wirth, R., Leal, I.R. 2019. Humanos e formigas utilizam as plantas que proliferaram com a fragmentação da mata atlântica nordestina. Ciência e Cultura, 71 (3), 58−61.

Câmara, T., Arnan, X., Barbosa, V., Wirth, R., Ianuzzi, L. and Leal, I.R. 2019. Disentangling the effects of foliar versus floral herbivory of leaf-cutting ants on the plant reproductive success of Miconia nervosa (Melastomataceae). Bulletin of Entomological Research, doi: 10.1017/S0007485319000294

Gerhold, V., Ribeiro, E. M. S., Santos, B. A., Sarapuu, J., Tabarelli, M., Wirth, R. and Leal, I. R. 2019. Phylogenetic signal in leaf-cutting ant diet in the fragmented Atlantic rain forest. Journal of Tropical Ecology. DOI: 10.1017/S0266467419000063

Siqueira, F. F. S., Ribeiro-Neto, J. D., Tabarelli, M., Andersen, A. N., Wirth, R. & Leal, I. R. 2018. Human disturbance promotes herbivory by leaf-cutting ants in the Caatinga dry forest. Biotropica. DOI: 10.1111/btp.12599

Dhodary, B., Schilg, M., Wirth, R. and Spiteller, D. S. 2018. Secondary metabolites from Escovopsis weberi and their role in attacking the garden fungus of leaf cutting ants. Chemistry - A European Journal, DOI:10.1002/chem.201706071.

Mueller, U. G., Ishak, H. D., Bruschi, S. M., Smith, C. C., Herman, J. J., Solomon, S. E., Mikheyev, A. S., Rabeling, C., Scott, J. J., Cooper, M., Rodrigues, A., Ortiz, A., Brandao, C. R. F., Lattke, J. E., Pagnocca, F. C., Rehner, S. A., Schultz, T. R., Vasconcelos, H. L., Adams, R. M. M., Bollazzi, M., Clark, R. M., Himler, A. G., LaPolla, J. S., Leal, I. R., Johnson, R. A., Roces, F., Sosa-Calvo, J., Wirth, R. and Bacci, M. 2017. Biogeography of Mutualistic Fungi Cultivated by Leafcutter Ants. Molecular Ecology DOI:10.1111/mec.14431

Silva, J. L. G., Silva, I. L. d. H., Ribeiro-Neto, J. D., Wirth, R. and Leal, I. R. 2017. Forest edge orientation influences leaf-cutting ant abundance and plant drought stress in the Brazilian Atlantic forest. Entomologia Experimentalis et Applicata DOI: 10.1111/afe.12268.

Siqueira, F.F.S., Ribeiro-Neto, J.D., Tabarelli, M., Andersen, A.N., Wirth, R., Leal, I.R. 2017. Leaf-cutting ant populations profit from human disturbances in tropical dry forest in Brazil. Journal of Tropical Ecology, doi:10.1017/S0266467417000311

Tabarelli, M., Siqueira, F. F. S., Backé, J., Wirth, R. & Leal, I. R. 2017. Ecology of leaf-cutting ants in human-modified landscapes. In: Oliveira, Paulo S. and Koptur, Suzanne (editors): Ant-Plant Interactions: Impacts of Humans on Terrestrial Ecosystems, p 73-90. Cambridge University Press

Mahé F., de Vargas C., Bass D., Czech L., Stamatakis A., Lara E., Singer D., Mayor J., Bunge J., Sernaker S., Siemensmeyer T., Trautmann I., Romac S., Berney C., Kozlov A., Mitchell E., Seppey C., Egge E., Lentendu G., Wirth R., Trueba G., Dunthorn M. 2017. Soil Protists in Three Neotropical Rainforests are Hyperdiverse and Dominated by Parasites. Nature Ecology & Evolution, doi:10.1038/s41559-017-0091

Baehner, K.W., Zweig, K.A., Leal, I.R., Wirth, R. 2017. Robustness of plant-insect herbivore interaction networks to climate change in a fragmented temperate forest landscape. Bulletin of Entomological Research: 1-10; DOI: 10.1017/S0007485317000062

Kelch, N.-S., Neves, F.S., Fernandes, G.W. & Wirth, R. 2016. Mechanisms driving galling success in a fragmented landscape: Synergy of habitat and top-down factors along temperate forest edges. PlosOne 11:e0157448

Corrêa, M.M., Silva, P.S.D., Wirth, R., Tabarelli, M. & Leal I.R. 2016. Foraging activity of leaf-cutting ants changes light availability and plant assemblage in Atlantic forest. Ecological Entomology,DOI: 10.1111/een.12312

Stephan, J. G., Wirth, R., Leal, I. R. & Meyer, S. T. 2015. Spatially heterogeneous nest-clearing behavior coincides with rain event in the leaf-cutting ant Atta cephalotes Linnaeus (Hymenoptera: Formicidae). Neotropical Entomology, DOI 10.1007/s13744-014-0267-0.

Leal, I. R., Wirth, R. & Tabarelli, M. 2014. The multiple impacts by leaf-cutting ants and their novel ecological role in human-modified neotropical forests. Biotropica 46: 516–528

Thiele, T., Kost, C., Roces, F. & Wirth, R. 2014. Foraging leaf-cutting ants learn to reject Vitis vinifera ssp. vinifera plants that emit herbivore-induced volatiles. Journal of Chemical Ecology 40: 617-620

Souza D.G., Santos B.A., Wirth R., Leal I.R. & Tabarelli M. 2013. Community-level patterns of insect herbivory in a fragmented Atlantic forest landscape. Environmental Entomology 42: 430-437

Meyer, S. T., Neubauer, M., Sayer, E. J., Leal, I. R., Tabarelli, M. & Wirth, R. 2013. Leaf-cutting ants as ecosystem engineers: topsoil and litter perturbations around Atta cephalotes nests reduce nutrient availability. Ecological Entomology, DOI: 10.1111/een.12043

Silva, P. S. D., Bieber, A. G., Knoch, T., Tabarelli, M., Leal, I. R. & Wirth, R. 2013. Foraging in highly dynamic environments: leaf-cutting ants adjust foraging trail networks to pioneer plant availability. Entomologia Experimentalis et Applicata DOI: 10.1111/eea.12050

Lakatos M., Wirth R., Spitzley P. & Lederer F. 2012. Der Biosphärenturm - ein innovatives Alleinstellungsmerkmal zur Baumkronenforschung. Ann. Sci. Rés. Bios. Trans. Vosges du Nord-Pfälzerwald 16: 118-129.

Ribeiro Neto, J. D., Pinho, B. X., Meyer, S. T., Wirth, R. & Leal, I. R. 2012. Drought stress drives intraspecific choice of food plants by Atta leaf-cutting ants. Entomologia Experimentalis et Applicata 144: 209-215.

Silva, P. S. D., Leal, I. R., Wirth, R. & Tabarelli, M. 2012. Spatial distribution and fruiting phenology of Protium heptaphyllum Aublet (March.) (Burceraceae) determine the design of the underground foraging system of Atta sexdens L. (Formicidae). Neotropical Entomology DOI 10.1007/s13744-012-0052-x.

Silva, P. S. D., Leal, I. R., Wirth, R., Melo, F. P. L. & Tabarelli, M. 2012. Leaf-cutting ants alter seedling assemblages across second-growth stands of Brazilian Atlantic forest. Journal of Tropical Ecology 28: 361–368.

Leal, I. R., Wirth, R. & Tabarelli, M. Formigas cortadeiras e a ambigüidade de suas relações com plantas. 2012. In: Del Claro, K. and Silingardi, H. M. T. Ecologia de Interações Plantas-Animais. Uma Abordagem Ecológico-Evolutiva,  p. 217-239. Technical Books Editora Ltda., Rio de Janeiro.

Colesie C., Scheu S., Green T.G., Weber B., Wirth R. & Büdel B. 2011. The advantage of growing on moss: facilitative effects on photosynthetic performance and growth in the cyanobacterial lichen Peltigera rufescens. Oecologia.  DOI: 10.1007/s00442-011-2224-5.

Dohm, C., Leal, I. R., Tabarelli, M., Meyer, S. T. & Wirth, R. 2011. Leaf-cutting ants proliferate in the Amazon: an expected response to forest edge? Journal of Tropical Ecology, 27: 645-649.

Kost, C., Tremmel, M. & Wirth, R. 2011. Do Leaf Cutting Ants Cut Undetected? Testing the Effect of Ant-Induced Plant Defences on Foraging Decisions in Atta colombica. PLoS ONE 6(7): e22340.

Schoenian, I., Spiteller, M., Ghaste, M., Wirth, R., Herz, H. & Spiteller, D. 2011. Chemical basis of the synergism and antagonism in microbial communities in the nests of leaf-cutting ants. Proceedings of the National Academy of Sciences doi:10.1073/pnas.1008441108.

Leal, I. R., Wirth, R. & Tabarelli, M. 2011. Dispersão de sementes por formigas cortadeiras. In: DellaLucia, T. M. C. Formigas cortadeiras. Da bioecologia ao manejo, p. 236-248. 2011. Editora da Universidade de Viçosa, Viçosa.

Meyer, S. T., Leal, I. R., Tabarelli, M. & Wirth, R. 2011. Ecosystem engineering by leaf-cutting ants: nests of Atta cephalotes drastically alter forest structure and microclimate. Ecological Entomology 36: 14-24.

Falcão, P. F., Pinto, S. R. R., Wirth, R. & Leal, I. R. 2011. Edge-induced narrowing of dietary diversity in leaf-cutting ants. Bulletin of Entomological Research 101: 305-311.

Bieber, A. G. D., Oliveira, M. A., Wirth, R., Tabarelli, M. & Leal, I. R. 2011. Do abandoned nests of leaf-cutting ants enhance plant recruitment in the Atlantic forest? Austral Ecology 36: 220-232.

Meyer, S. T., Leal, I. R., Tabarelli, M. & Wirth, R. 2011. Performance and fate of tree seedlings on and around nests of the leaf-cutting ant Atta cephalotes: Ecological filters in a fragmented forest. Austral Ecology 36: 779–790.

Corrêa, M., Silva, P., Wirth, R., Tabarelli, M. & Leal, I. 2010. How leaf-cutting ants impact forests: drastic nest effects on light environment and plant assemblages. Oecologia 162: 103–115.

Wirth, R. 2009. Book review: The tropical mountain forest - patterns and processes in a biodiversity hotspot, S.R. Gradstein, J. Homeier, D. Gansert (Eds). Universitätsverlag Göttingen, Göttingen (2008). 217 pp. 38,00 €, ISBN: 978-3-940344-22-9. In: Basic and Applied Ecology 10: 684-685.

Meyer, S. T., Leal, I.R., Wirth, R. 2009. Persisting hyper-abundance of keystone herbivores (Atta spp.) at the edge of an old Brazilian Atlantic Forest fragment. Biotropica 41: 711-716.

Haeder S., Wirth R., Herz H., Spiteller, D. 2009. Candicidin-producing Streptomyces support leaf-cutting ants to protect their fungus garden against the pathogenic fungus Escovopsis. PNAS 106: 4742-4746.

Silva, P.S.D., Bieber, A.G.D., Leal, I.R., Wirth R., Tabarelli, M. 2009. Decreasing abundance of leaf-cutting ants across a chronosequence of advancing Atlantic forest regeneration. Journal of Tropical Ecology 25:1–5.

Almeida, W. R., Wirth, R., Leal, I. R. 2008. Edge-mediated reduction of phorid parasitism on leaf-cutting ants in a Brazilian Atlantic forest. Entomologia Experimentalis et Applicata, 129: 251-257.

Wirth, R., Meyer, S.T., Leal, I.R. and Tabarelli, M. 2008. Plant Herbivore Interactions at the Forest Edge. In: Lüttge, U., Beyschlag, W. and Murata, J. (eds). Progress in Botany 69: 423-448. Springer, Berlin Heidelberg.

Silva, P. D., Leal, I.R., Wirth, R., Tabarelli, M. 2007. Harvesting of Protium heptaphyllum Aublet (March.) seeds (Burseraceae) by the leaf-cutting ant Atta sexdens L. promotes seed aggregation and seedling mortality". Revista Brasileira de Botânica 30: 553-560.

Wirth, R., Meyer, S.T., Almeida, W.R., Araújo Jr, M.V., Barbosa, V.S., Leal, I.R. 2007. Increasing densities of leaf-cutting ants (Atta spp.) with proximity to the edge in a Brazilian Atlantic forest. Journal of Tropical Ecology 23: 501-505.

Kost, Ch., Lakatos, T., Arendholz, W-R., Redenbach, M. & Wirth, R. 2007. Unspecific association between filamentous bacteria and fungus-growing ants. Naturwissenschaften, DOI 10.1007/s00114-007-0262-y.

Urbas, P., Araújo Jr., M., Leal, I.R. Wirth, R. 2007. Cutting More from cut forests - edge effects on foraging and herbivory of leaf-cutting ants. Biotropica 39: 489-495.

Leal, I. R., Wirth, R. & Tabarelli, M. 2007. Seed dispersal by ants in the semi-arid Caatinga of northeast Brazil. Annals of Botany 99: 885-894.

Meyer, S., Roces, F., Wirth, R. 2006. Selecting the drought stressed: Effects of plant stress on intraspecific and within-plant herbivory patterns of the leaf-cutting ant Atta colombica. Functional Ecology 20:973-981.

Leal, I.R, Fischer, E., Kost, Ch., Tabarelli, M., Wirth, R. 2006. Ant protection against herbivores and nectar thieves in Passiflora coccinea flowers. Ecoscience: 13: 431-438.

Corrêa, M.M., Bieber, A.G.D, Wirth, R. & Leal, I.R. 2005 Occurrence of Atta cephalotes (L.) (Hymenoptera: Formicidae) in Alagoas, Northeastern Brazil. Neotropical Entomology, 34 : 695-698.

Kost C., Gama de Oliveira E., Knoch T. & Wirth R. 2005. Temporal and spatial patterns, plasticity, and ontogeny of foraging trails in leaf-cutting ants. Journal of Tropical Ecology, 21: 677 – 688.

Berger, J. & Wirth, R. 2004. Predation-mediated Mortality of Early Life Stages: A Field Experiment with Nymphs of an Herbivorous Stick Insect (Metriophasma diocles). Biotropica, 36 :424-428.

Wirth, R., Beyschlag, W., Ryel, R., Herz, H. & Hölldobler, B. 2003. The herbivory of leaf-cutting ants. A case study on Atta colombica in the tropical rainforest of Panama. Accepted by Ecological Studies, Springer Verlag Berlin, Heidelberg, New York. Cover and abstract

Wirth, R., Weber, B. & Ryel, R. 2001. Spatial and temporal variability of canopy structure in a tropical moist forest. Acta Oecologia 22: 235-244.

Wirth, R. & Leal, I. 2001. Rainfall affects temporal variability of ant protection in Passiflora coccinea. Ecoscience, 450-453.

Wirth, R., Kost, C., Schmidt, S. 2001. Costs of trail construction and maintenance may affect the pattern of herbivory in leaf-cutting ants. In: G. Zotz, Ch. Körner (eds.) The functional importance of biodiversity. Verhandlungen der Gesellschaft für Ökologie 31: 315.

Berger, J., Wirth, R. 2001. Physical antiherbivore defense does not determine preference of a specialist stick insect. In: G. Zotz, Ch. Körner (eds.) The functional importance of biodiversity. Verhandlungen der Gesellschaft für Ökologie 31: 311.

Berger, J., Wirth, R. 2001. Diversity and diet breadth of stick insects (Phasmatodea) in a Panamanian semi-deciduous rainforest). In: G. Zotz, Ch. Körner (eds.) The functional importance of biodiversity. Verhandlungen der Gesellschaft für Ökologie 31: 338.

Wirth, R., Kirsch, H. & Büdel, B. 2001. Verbreitungsmuster und Dynamik der Wiederausbreitung von Bartflechten der Gattungen Usnea und Bryoria im Spessart. Hoppea, Denkschr. Regensb. Bot. Ges, 62: 411-436.

Wirth, R. 1999. Blattschneiderameisen und Pflanzen: Wer kontrolliert wen? Ausmaß und Muster der Herbivorie von Blattschneiderameisen. Pg. 203-215 in Beyschlag, W. & Steinlein, T. (eds.). Ökophysiologie pflanzlicher Interaktionen. Bielefelder Ökologische Beiträge 14: 203-215.

Wirth, R., Weigelt, A., Herz, H., Hölldobler, B. & Beyschlag, W. 1998. Differential herbivory by leaf-cutting ants and consequences for light availability within tropical forest vegetation. In Dalitz, H., Haverkamp, M., Homeier, J. & Breckle, S.W. (eds.). GTÖ 98. Kurzbeiträge zur Tropenökologie. Bielefelder Ökologische Beiträge 12.

Dalling, J. W. & Wirth, R. 1998. Dispersal of Miconia argentea seeds by the leaf-cutting ant Atta colombica. Journal of Tropical Ecology 14: 705-710.

Herz, H., Wirth, R., Hölldobler, B. & Beyschlag, W. 1998. High rate of leaf-cutter ant colony movements in a tropical forest. In: Dalitz H, Haverkamp M, Homeier J, Breckle SW (eds.). GTÖ 98. Kurzbeiträge zur Tropenökologie. Bielefelder Ökologische Beiträge 12.

Wirth, R., Weigelt, A., Herz, H., Hölldobler, B. & Beyschlag, W. 1998. Differential herbivory by leaf-cutting ants and consequences for light availability within tropical forest vegetation. In Dalitz, H., Haverkamp, M., Homeier, J. & Breckle, S.W. (eds.). GTÖ 98. Kurzbeiträge zur Tropenökologie. Bielefelder Ökologische Beiträge 12.

Wirth, R., Beyschlag, W., Ryel, R. J. & Hölldobler, B. 1997. Annual foraging of the leaf-cutting ant Atta colombica in a semideciduous rain forest in Panama. Journal of Tropical Ecology 13: 741-757.

Wirth, R., & Büdel B. 1994. Das Auftreten von Pycnidien bei fruchtenden und sterilen Formen von Usnea florida (L.) Hoff. (Lecanorales, Usneaceae). Nova Hedwigia 58: 13-20.