Variable Success of Biological Control Agents for Lythrum salicaria in Minnesota Wetlands: Understanding Landscape Patterns in Plant Evolution and Management Efficacy Evolution of Increased Competitive Ability (EICA) Hypothesis • Specialist herbivores removed Introduced Populations • Cost but no benefit of defense Time (Blossey & Nötzold 1995; Reviewed by Bossdorf et al., 2005) Evolution of Increased Competitive Ability (EICA) Hypothesis • Specialist herbivores removed Introduced Populations • Cost but no benefit of defense • Plants evolve to allocate resources to vigor (Blossey & Nötzold 1995; Reviewed by Bossdorf et al., 2005) Evolution of Increased vigor from loss of herbivore defense Time L. Salicaria Evidence for Evolution of Increased Competitive Ability L. Salicaria Evidence for Evolution of Increased Competitive Ability • Enemy Release – Specialist herbivores (Hight, 1990) L. Salicaria Evidence for Evolution of Increased Competitive Ability • Enemy Release – Specialist herbivores (Hight, 1990) • Evolution of Increased Competitive Ability – Decreased herbivore resistance (Blossey & Nötzold, 1995; Blossey & Kamil, 1996) – Increased Vigor (Willis et al., 1999; Blossey & Nötzold, 1995; Blossey & Kamil, 1996; Willis & Blossey, 1999; Bastlova & Květ, 2002; Chun et al., 2007) Lythrum salicaria Biocontrol • In 1992 three species were introduced • 2,150 Beetles were released in Minnesota at two sites (Malecki, R. et al 1993; MN DNR, 2009) Lythrum salicaria Biocontrol • In 1992 three species were introduced • 2,150 Beetles were released in Minnesota at two sites • More than 9 million insects released in Minnesota (Malecki, R. et al 1993; MN DNR, 2009) Invasion from Europe; release from specialist herbivores ~200 Years ago Evolutionary increased vigor from loss of herbivore defense Introduction of biocontrol agents 1992 Invasion from Europe; release from specialist herbivores ~200 Years ago Evolutionary increased vigor from loss of herbivore defense Introduction of biocontrol agents 1992 Evolutionary response to selective pressure of specialist herbivores? ? Invasion from Europe; release from specialist herbivores ~200 Years ago Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents ? “Historically Low Herbivory (LH)” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents “Historically High Herbivory (HH)” ? “Historically Low Herbivory (LH)” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents Evolutionary divergence in response to selective pressure of specialist herbivores? “Historically High Herbivory (HH)” ? Experimental Garden Design ML 1 ML 2 Wetland 1 Wetland 2 Wetland 3 Wetland 4 Wetland 5 Harvest 1st year Wetland 6 = LH = HH Harvest 2nd year ML 3 ML 4 ML 5 ML 6 H e = LH C = HH W in Pi g' s ke nd on a Ey e la Po e y sh te r M ar Ce n irc l tte r al l's od g Po D (g ( 95%CI) Weight Total L. salicaria DryGarden Data (2009) Experimental Lythrum salicaria Vigor 1st Year Average Total Weight (g) 150 100 50 0 Lythrum salicaria Vigor 1st Year • Stem Weight 150 < • Inflorescence Weight 100 < • Stem Height 50 < on a W in Ey e Pi g' s ke e irc l C y tte r Po H al l's = LH la Po sh M ar te r Ce n e od g nd 0 D (g ( 95%CI) Weight Total L. salicaria DryGarden Data (2009) Experimental Average Total Weight (g) = HH (Nested ANOVA Models, p < 0.05) Lythrum salicaria Vigor 2nd Year • Litter Weight < • Stem Weight < • Stem Weight < • Inflorescence Weight < • Inflorescence Weight < • Root Weight • Stem Height < < • Total Plant Weight < = LH = HH (Nested ANOVA Models, p < 0.05) • Stem Height < Lythrum salicaria Herbivore Defense • Tannins- defense against specialist herbivores (Muller-Scharer 2004) • L. salicaria leaf dry weight 24% tannins (Rauha et al., 2000) Lythrum salicaria Herbivore Defense • Tannins- defense against specialist herbivores (Muller-Scharer 2004) • L. salicaria leaf dry weight 24% tannins (Rauha et al., 2000) = LH = HH Tannins (mg/g ( 95%C) 65 60 55 50 45 40 HLH Significant effect of population type (Nested ANOVA F= 4.095 p = 0.044) HHH Experimental Garden Design ML 1 ML 2 Wetland 1 Wetland 2 Wetland 3 Wetland 4 Wetland 5 Harvest 1st year Wetland 6 = LH = HH Harvest 2nd year ML 3 ML 4 ML 5 ML 6 Experimental Garden Design ML 1 Wetland 1 Wetland 2 Wetland 3 Wetland 4 Wetland 5 Harvest 1st year Wetland 6 = LH = HH Harvest 2nd year ML 2 ML 3 ML 4 ML 5 ML 6 Lythrum salicaria Herbivore Tolerance • HH population exhibited compensatory growth L. salicaria Total Weight (g) 300 AC C C 250 AB 200 B B 150 100 50 0 HLH Alone = LH HLH w/Typha HLH w/Biocontrol = HH (Nested ANOVA Models Tukey HSD, p < 0.05) HHH Alone HHH w/Typha HHH w/Biocontrol Lythrum salicaria Herbivore Tolerance • Herbivory increased HH population reproduction 9 AC Inflorescence Weight (g) 8 7 C C AB 6 B B HLH w/Typha HLH w/Biocontrol 5 4 3 2 1 0 HLH Alone = LH = HH (Nested ANOVA Models Tukey HSD, p < 0.05) HHH Alone HHH w/Typha HHH w/Biocontrol “Historically Low Herbivory” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents Evolutionary divergence in response to selective pressure of specialist herbivores? “Historically High Herbivory” ? “Historically Low Herbivory” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents Evolutionary divergence in response to selective pressure of specialist herbivores? “Historically High Herbivory” “Historically Low Herbivory” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents Does this evolutionary divergence in plant traits affect Galrecuella behavior? “Historically High Herbivory” “Historically Low Herbivory” Invasion from Europe; release from specialist herbivores ~200 Years ago 1992 Introduction of biocontrol agents How will the presence of biocontrol agents drive evolution of L. salicaria in the future? “Historically High Herbivory” Acknowledgements Thank you to: Funding: Jeannine Cavender-Bares Will Menzel Ruth Shaw Diane Larson Mia Howard Neil Anderson Steven Lee Luke Skinner Jennie Sirota Alyssa Bernardo Jan Kvet Dasa Bastlova Rodrigo Gonzalez Chauvet Lukas Johnson Leon & Naomi Quiram 30 construction volunteers Dayton Wilkie Fellowship Fund MN-Department of Natural Resources Carolyn Crosby Fellowship University of Minnesota Graduate School MNCenter for Community Genetics Plant Traits Seminar Small Grants Rothman Fellowship Fund Dept. of Ecology, Evolution and Behavior Dayton Fellowship Fund Pletcher Graduate Fellowship Alexander Dubcek Fund NSF ISG-IGERT Traineeship Galerucella spp. Choice Trials • Grew plants from HH & LH populations • Collected Galerucella from HH & LH populations • Choice tests Occupancy and behavior Feeding preference Egg laying preference Galerucella spp. choice Frequency of Occupation • No preference in – feeding – oviposition Laying Eggs 0.1% Mating 9.3% Eating 12.0% Still 67.8% Other 0.4% Eating 12.8% Walking 8.1% 18 16 14 12 10 8 6 4 2 0 LH Low HH High Walking 10.8% Still 78.8% (Paired t-test; p= 0.01570, n= 77) Site Location/Herbivory History “Historically Low Herbivory” “Historically High Herbivory” Site Name Dodge Nature Center 1998 1999 2000 2001 F D D D Hall's Marsh D Pottery Pond 2002 2003 2004 D C C C D C C 2007 D C D A Winona B A A A A A A A Circle Lake A A A A A A A A Pig's Eye* D F D D A A= widespread >80-100% defoliation of entire wetland B= 50-90% defoliation generally widespread C= damage from insects near release points D= No damage, few insects, not established F= No insects or damage present * The data was collected on a transect that did not run thought the area of beetle control. ** Locations for HHH and HLH sites taken from DNR biological control agent release records. For native sites location was estimated from a Shocart map 215, ISBN: 978-80-7224-267-2. A Invertebrates on Lythrum salicaria in the US http://www.flickr.com/groups/lifeontheloosestrife Minnesota Biocontrol Purple loosestrife infestation in Minnesota Data from: MN DNR Insect release sites in Minnesota
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