title: How slug herbivory of juvenile hybrid willows alters chemistry, growth and subsequent susceptibility to diverse plant enemies.
abstract: OBJECTIVE
  Selective feeding by herbivores, especially at the seedling or juvenile phase, has the potential to change plant traits and ultimately the susceptibility of surviving plants to other enemies. Moreover, since hybridization is important to speciation and can lead to introgression of traits between plant species, differential feeding (herbivore-induced mortality) can influence the expression of resistance traits of hybrids and ultimately determine the consequences of hybridization. While it would be expected that herbivore-induced mortality would lead to greater resistance, there may be trade-offs whereby resistance to one herbivore increases susceptibility to others. The hypothesis was tested that the exotic slug, Arion subfuscus, causes non-random survival of hybrid willows and alters plant: (1) susceptibility to slugs; (2) secondary and nutritional chemistry, and growth; and (3) susceptibility to other phytophages.
  
  
  METHODS
  Two populations of plants, control and selected, were created by placing trays of juvenile willows in the field and allowing slugs access to only some. When ≤10 individuals/tray remained (approx. 85 % mortality), 'selected' and undamaged 'control' trays were returned to a common area. Traits of these populations were then examined in year 1 and in subsequent years.
  
  
  RESULTS
  The selected population was less palatable to slugs. Surprisingly, foliar concentrations of putative defence traits (phenolic glycosides and tannins) did not differ between treatments, but the selected population had higher foliar nitrogen and protein, lower carbon to nitrogen ratio and greater above-ground biomass, indicating that vigorously growing plants were inherently more resistant to slugs. Interestingly, selected plants were more susceptible to three phytophages: an indigenous pathogen (Melampsora epitea), a native herbivorous beetle (Chrysomela knabi) and an exotic willow leaf beetle (Plagiodera versicolora).
  
  
  CONCLUSIONS
  This exotic slug changed the population structure of F2 hybrid willows in unanticipated ways. Defence expression remained unchanged, while nutritional and growth traits changed. These changes caused plants to be more susceptible to other plant enemies. Other exotic herbivore species are anticipated to have similar direct and indirect effects on native plant populations.
authors.affiliation: Department of Biology, Tufts University, Medford, MA 02155, USA. colin.orians@tufts.edu
authors.forename: Colin M
authors.initials: CM
authors.lastname: Orians
authors.forename: Robert S
authors.initials: RS
authors.lastname: Fritz
authors.forename: Cris G
authors.initials: CG
authors.lastname: Hochwender
authors.forename: Benedicte R
authors.initials: BR
authors.lastname: Albrectsen
authors.forename: Mary Ellen
authors.initials: ME
authors.lastname: Czesak
mesh_terms: D000818:Animals; D049851:Gastropoda / Q000502:physiology*; D006027:Glycosides / Q000378:metabolism; D060434:Herbivory; D010636:Phenols / Q000378:metabolism; D032108:Salix / Q000502:physiology*; D012641:Selection, Genetic; D013634:Tannins / Q000378:metabolism
chemical_list: D006027:Glycosides; D010636:Phenols; D013634:Tannins
keywords: Arion subfuscus; Exotic species; Salix; herbivory; hybridization; slug; trade-offs; willow