, 150:421 | Cite as

On the elusiveness of enemy-free space: spatial, temporal, and host-plant-related variation in parasitoid attack rates on three gallmakers of goldenrods

  • Stephen B. Heard
  • John O. Stireman III
  • John D. Nason
  • Graham H. Cox
  • Christopher R. Kolacz
  • Jonathan M. Brown
Plant Animal Interactions


Host shifting by phytophagous insects may play an important role in generating insect diversity by initiating host-race formation and speciation. Models of the host shifting process often invoke reduced rates of natural enemy attack on a novel host in order to balance the maladaptation expected following the shift. Such “enemy-free space” has been documented for some insects, at some times and places, but few studies have assessed the occurrence of enemy-free space across years, among sites, or among insect species. We measured parasitoid attack rates on three insect herbivores of two goldenrods (Solidago altissima L. and Solidago gigantea Ait.), with data from multiple sites and multiple years for each herbivore. For each insect herbivore, there were times and sites at which parasitoid attack rates differed strongly and significantly between host plants (that is, enemy-free space existed on one host plant or the other). However, the extent and even the direction of the attack-rate difference varied strongly among sites and even among years at the same site. There was no evidence of consistent enemy-free space for any herbivore on either host plant. Our data suggest that enemy-free space, like many ecological and evolutionary forces, is likely to operate as a geographic and temporal mosaic, and that conceptual models of host shifting that include enemy-free space as a consequence of host novelty are likely too simple.


Host shift Enemy-free space Parasitoid Phytophagous insect 



For permission to collect goldenrod galls, we thank Gerald and Bonnie Kragt, the City of Fredericton, the Iowa Department of Natural Resources, the Toronto and Region Conservation Authority, and the U.S. Fish and Wildlife Service. Kevin Day, William Godsoe, Kristie Heard, Michelle LeBlanc, Trina Nowack, Katie Richardson, Kristie Richardson, Julie Seehawer, David Smith, Paige Smith, Kristina Tarvin, Susan Timmons, Fred Williams, Dennis Wong, and three generations of Terpstras and Hanenbergs assisted in field and lab. Comments by Kristie Heard, John Reeve, and two anonymous reviewers vastly improved the manuscript. This research was funded by the Natural Sciences and Engineering Research Council (Canada; Discovery Grant to SBH) and by the National Science Foundation (USA; grants DEB 0707752 to SBH and DEB 0107938 to JDN).

Supplementary material

442_2006_529_MOESM1_ESM.doc (72 kb)
Supplementary material


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Copyright information

© Springer-Verlag 2006

Authors and Affiliations

  • Stephen B. Heard
    • 1
  • John O. Stireman III
    • 2
    • 3
  • John D. Nason
    • 2
  • Graham H. Cox
    • 1
  • Christopher R. Kolacz
    • 1
  • Jonathan M. Brown
    • 4
  1. 1.Department of BiologyUniversity of New BrunswickFrederictonCanada
  2. 2.Department of Ecology, Evolution, and Organismal BiologyIowa State UniversityAmesUSA
  3. 3.Department of Biological SciencesWright State UniversityDaytonUSA
  4. 4.Department of BiologyGrinnell CollegeGrinnellUSA

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