title: Rapid evolution of virulence leading to host extinction under host-parasite coevolution.
abstract: BACKGROUND
  Host-parasite coevolution is predicted to result in changes in the virulence of the parasite in order to maximise its reproductive success and transmission potential, either via direct host-to-host transfer or through the environment. The majority of coevolution experiments, however, do not allow for environmental transmission or persistence of long lived parasite stages, in spite of the fact that these may be critical for the evolutionary success of spore forming parasites under natural conditions. We carried out a coevolution experiment using the red flour beetle, Tribolium castaneum, and its natural microsporidian parasite, Paranosema whitei. Beetles and their environment, inclusive of spores released into it, were transferred from generation to generation. We additionally took a modelling approach to further assess the importance of transmissive parasite stages on virulence evolution.
  
  
  RESULTS
  In all parasite treatments of the experiment, coevolution resulted in extinction of the host population, with a pronounced increase in virulence being seen. Our modelling approach highlighted the presence of environmental transmissive parasite stages as being critical to the trajectory of virulence evolution in this system.
  
  
  CONCLUSIONS
  The extinction of host populations was unexpected, particularly as parasite virulence is often seen to decrease in host-parasite coevolution. This, in combination with the increase in virulence and results obtained from the model, suggest that the inclusion of transmissive parasite stages is important to improving our understanding of virulence evolution.
authors.affiliation: Evolutionary Ecology and Genetics, Zoological Institute, Christian-Albrechts-Universitaet zu Kiel, Am Botanischen Garten 1-9, Kiel, 24118, Germany. charlotte.rafaluk@gmail.com.|Department of Zoology, University of Oxford, The Tinburgen Building, South Parks Road, Oxford, OX1 3PS, UK. charlotte.rafaluk@gmail.com.|Institute for Phytopathology and Applied Zoology, University of Giessen, Heinrich-Buff-Ring 26-32, Gießen, D-35392, Germany. charlotte.rafaluk@gmail.com.
authors.forename: Charlotte
authors.initials: C
authors.lastname: Rafaluk
authors.affiliation: Evolutionary Ecology and Genetics, Zoological Institute, Christian-Albrechts-Universitaet zu Kiel, Am Botanischen Garten 1-9, Kiel, 24118, Germany. gildenhard@mpiib-berlin.mpg.de.|Department of Vector Biology, Max-Planck-Institut für Infektionsbiologie, Campus Charité Mitte, Charitéplatz 1, Berlin, 10117, Germany. gildenhard@mpiib-berlin.mpg.de.
authors.forename: Markus
authors.initials: M
authors.lastname: Gildenhard
authors.affiliation: Evolutionary Ecology and Genetics, Zoological Institute, Christian-Albrechts-Universitaet zu Kiel, Am Botanischen Garten 1-9, Kiel, 24118, Germany. andreas.mitschke@agrar.uni-giessen.de.|Institute for Phytopathology and Applied Zoology, University of Giessen, Heinrich-Buff-Ring 26-32, Gießen, D-35392, Germany. andreas.mitschke@agrar.uni-giessen.de.
authors.forename: Andreas
authors.initials: A
authors.lastname: Mitschke
authors.affiliation: Westfälische Wilhelms-Universität Münster, Institut für Evolution und Biodiversität, Hüfferstr. 1, 48149, Münster, Germany. Arndt.Telschow@uni-muenster.de.
authors.forename: Arndt
authors.initials: A
authors.lastname: Telschow
authors.affiliation: Evolutionary Ecology and Genetics, Zoological Institute, Christian-Albrechts-Universitaet zu Kiel, Am Botanischen Garten 1-9, Kiel, 24118, Germany. hschulenburg@zoologie.uni-kiel.de.
authors.forename: Hinrich
authors.initials: H
authors.lastname: Schulenburg
authors.affiliation: Evolutionary Ecology and Genetics, Zoological Institute, Christian-Albrechts-Universitaet zu Kiel, Am Botanischen Garten 1-9, Kiel, 24118, Germany. Gerrit.Joop@agrar.uni-giessen.de.|Institute for Phytopathology and Applied Zoology, University of Giessen, Heinrich-Buff-Ring 26-32, Gießen, D-35392, Germany. Gerrit.Joop@agrar.uni-giessen.de.
authors.forename: Gerrit
authors.initials: G
authors.lastname: Joop
mesh_terms: D000818:Animals; D005075:Biological Evolution; D004777:Environment; D006790:Host-Parasite Interactions; D016818:Nosema / Q000235:genetics* / Q000472:pathogenicity* / Q000502:physiology; D014231:Tribolium / Q000235:genetics* / Q000469:parasitology*; D014774:Virulence