Final Report Illlinois-Indiana Sea Grant June 30, 2011

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1 Ecological genetics of Phragmites australis invasion in southern Lake Michigan coastal habitats (Seed Project) Final Report Illlinois-Indiana Sea Grant June 30, 2011 Investigators: Dr. Daniel Larkin Plant Science and Conservation Chicago Botanic Garden 1000 Lake Cook Road Glencoe, IL Dr. Jeremie Fant Plant Science and Conservation Chicago Botanic Garden 1000 Lake Cook Road Glencoe, IL

2 Project summary Under this award, the project team investigated the genetics of populations of Phragmites australis (common reed) throughout the Lake Michigan coast of Illinois and Indiana. In a parallel study, the environmental correlates and plant-community consequences of Phragmites invasion in coastal and inland wetlands in northeast Illinois and northwest Indiana were observed. A total of 71 patches of Phragmites distributed across 17 sites (Figure 1) were sampled, analyzing the genetics of 450 individual plants. The exotic subspecies, P. australis ssp. Australis, occurred with greater frequency than the native subspecies, P. australis ssp. americanus (Table 1, subspecies determined by RFLP fingerprinting). Exotic genotypes recurred less frequently among different sites than native genotypes, which is consistent with a more recent colonization. Despite Phragmites expansion being widely thought of as a function of its ability to spread clonally, the high genetic diversity within and between patches is consistent with colonization through seed rather than stem fragments (Figure 2, Figure 3). An analysis of microsatellite data using Structure v2.0 allowed for comparison of the genetic structure of populations sampled with reference populations analyzed in publications by Saltonstall. Native Phragmites were found to be a distinct, homogenous group, of the same genetic origins as reference populations (Figure 4). However, exotic Phragmites appeared to comprise two distinct genetic groups, one of which was not well-represented in reference collections. This suggests that the Illinois-Indiana region may be subject to invasion by a distinct exotic strain. In parallel to the genetic work described above, an environmental and ecological sampling was performed to elucidate the behavior of native vs. exotic Phragmites. Attributes such as the amount of bare ground, water cover, water depth, soil moisture, and salinity were not important in explaining the occurrence of uninvaded patches versus native-phragmites patches versus exotic-phragmites patches. Inorganic nitrogen availability (nitrate + ammonium) significantly differed among patches of different invasion status, but not in a consistent manner and without indication that increasing nitrogen was associated with more aggressive Phragmites growth (Figure 5, Figure 6). In contrast, phosphorus availability was higher in invaded areas, and exotic Phragmites in particular showed a strong positive growth response to phosphorus availability (Figure 7, Figure 8). Exotic but not native Phragmites were associated with strong local losses of plant diversity (Figure 9). Based on these seed results, the following conclusions were drawn for management of coastal habitats colonized by Phragmites: Exotic and native Phragmites are quite different genetically and ecologically despite being conspecifics. No evidence of the native subspecies behaving invasively was found in the study. Wetland managers should prioritize control efforts on the exotic subspecies. Other studies have found evidence of hybridization between Phragmites subspecies. Were hybridization to occur in Illinois-Indiana coastal habitats, Phragmites could pose an even greater ecological threat due to hybrid vigor. Further investigation is needed on Phragmites dispersal dynamics and rate of spread, the effects of land-use on invasion risk, and the effectiveness of current efforts to control Phragmites and restore diverse native plant communities in degraded coastal systems. Larkin and Fant 2

3 Students supported and degrees conferred Research conducted as part of this award contributed to research experiences for one graduate, one post-bachelors, and four undergraduate students: This award supported the master's thesis research of Amy Price, who will receive her M.S. in Plant Biology and Conservation from Northwestern University (defending July 2011). Research Assistant Joseph Boyer performed field and laboratory work under this award, helping him secure a position as a paid conservation intern with the Bureau of Land Management in Richland, Utah. Field work was conducted by undergraduate researcher Cat Collins (DePaul University). Lab work was conducted by undergraduate researchers Adewale Adeoba and David Ford (Loyola University Chicago) and Dara Wise (Benedictine University, accepted to medical school for fall 2011). Publications and presentations Publications Price, A.L. In prep (Jul 2011 completion). Genetics and ecology of Phragmites australis invasion in the upper Midwest. Master's Thesis. Northwestern University, Evanston, IL. Price, A.L., J.B. Fant, and D.J. Larkin. In prep. Comparative ecology of native vs. exotic Phragmites australis (common reed) in Chicago-area wetlands. For submission to Wetlands. Fant, J.B., A.L. Price, and D.J. Larkin. In prep. Genetic structure and reproduction of native and exotic Phragmites australis. For submission to Conservation Genetics. Presentations Larkin, D.J., A.L. Price, and J.B. Fant When the species concept and land management collide: Phragmites australis invasion in the Chicago region. The Chicago Plant Science Symposium. Chicago, IL. Apr 16, Larkin, D.J., A.L. Price, and J.B. Fant Ecology and genetics of Phragmites australis invasion in the Chicago region. Presentation to the Chicago Wilderness Natural Resources Management Team. Chicago, IL. Apr 25, Larkin and Fant 3

4 Tables and Figures Table 1. Occurrence of unique native and exotic genotypes of Phragmites in sampled patches. Native Exotic Genotypes Patches found in 1 74% 98% 2 21% 2% 3 5% 0% 4 0% 0% Sites found in 1 100% 100% 2 0% 0% IL Beach S.P. Lake Total: 71 Phragmites patches, 17 sites Intensive sampling (6 sites) (genetic, environmental, ecological) Genetic only Kane DuPage Cook Calumet IN Dunes N.L. Figure 1. Locations of Phragmites australis populations sampled as part of this study. 2 Larkin and Fant 4

5 Proportion of genotypes per patch all individuals unique (G-1)/(N-1) all individuals identical IBSP-1 IBSP-2 IBSP-3 IBSP-4 IBSP-5 IBSP-6 IBSP-7 IBSP-8 IBSP-9 IBSP-10 Illinois Beach State Park IBSP-11 IBSP-12 IDNL-1 IDNL-2 IDNL-3 IDNL-4.1 IDNL-4.2 IDNL-4.3 IDNL-4.4 IDNL-5 IDNL-6 Indiana Dunes N.L. IDNL-7 WCP1 WCP2 W. Chi. Prairie WG1 WW1 WW2 WW3 WW4 Evanston Figure 2. Patch-level genetic diversity for subset of sites sampled in this study. Blue and orange bars indicate representatives of the native and exotic subspecies, respectively. Results indicate that sexual reproduction is an important factor within patches, despite Phragmites being a clonal species. WW5 ssp. americanus (native) among sites 15% ssp. australis (exotic) among sites 16% within patches 55% among patches 30% within patches 48% among patches 36% Figure 3. AMOVA analysis showing the distribution of genetic diversity among different population levels. Larkin and Fant 5

6 100% 80% 60% 40% native exotic 1 exotic 2 20% 0% 1 Saltonstall IBSP IDNL Evanston W. Chi. (NU) Prairie Figure 4. Results of a structure analysis of microsatellite data. Saltonstall data used for reference populations. Calumet Prairie Dick Young Warrenville Grove P = TIN (NH3+ NOX) (μg g 1 dry soil) Uninvaded PHRAUS PHRAUS ssp. americanus ssp. australis Phragmites status Figure 5. Differences in sediment inorganic nitrogen concentration in uninvaded, native-phragmites, and exotic-phragmites patches. Error bars are ± 1. s.e. Larkin and Fant 6

7 80 No relationship (both ssp.) uninvaded ssp. americanus ssp. australis 60 Phragmites density (stems m 2 ) TIN (NH 3 + NO X ) log(μg g 1 dry soil) Figure 6. There was no increase in Phragmites stem density with increasing inorganic nitrogen availability P = (μg g 1 dry soil) PO Uninvaded PHRAUS PHRAUS ssp. americanus ssp. australis Phragmites status Figure 7. Differences in sediment orthophosphate concentration in uninvaded, native-phragmites, and exotic-phragmites patches. Error bars are ± 1. s.e. Larkin and Fant 7

8 60 50 uninvaded ssp. americanus ssp. australis australis americanus ANCOVA: P = 0.05 Phragmites density (stems m 2 ) PO 4 log(μg g 1 dry soil + 1) Figure 8. Phragmites stem density increased with orthophosphate availability, particularly for the exotic subspecies. 2.5 australis americanus ANCOVA: P < uninvaded ssp. americanus ssp. australis Shannon-Wiener diversity (H') r 2 = 0.06 r 2 = Phragmites density (stems m 2 ) Figure 9. Non-Phragmites plant diversity was weakly positively correlated with stem density of native Phragmites but strongly, negatively correlated with increasing density of exotic Phragmites. Larkin and Fant 8

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