A Comparative Assessment of Seedling Survival and Biomass Accumulation for Fourteen Wetland Plant Species Grown Under Minor Water-Depth Differences
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WETLANDS, Vol. 25, No. 3, September 2005, pp. 520–530 ᭧ 2005, The Society of Wetland Scientists A COMPARATIVE ASSESSMENT OF SEEDLING SURVIVAL AND BIOMASS ACCUMULATION FOR FOURTEEN WETLAND PLANT SPECIES GROWN UNDER MINOR WATER-DEPTH DIFFERENCES Lauchlan H. Fraser1,2 and Jason P. Karnezis1 1Department of Biology University of Akron Akron, Ohio, USA 44325-3908 2Present address: Department of Natural Resource Science Thompson Rivers University Kamloops, British Columbia, Canada V2C 5N3 E-mail: [email protected] Abstract: The purpose of this experiment was to investigate the survival and biomass accumulation of wetland plant species under different water depths in controlled microcosms. In the greenhouse, two-week- old seedlings were randomly assigned to one of seven water-depth treatments (Ϫ6, Ϫ4, Ϫ2, 0, ϩ2, ϩ4, and ϩ6 cm relative to the soil surface) and allowed to grow for six months. Species included five perennial sedges, four perennial and one annual grasses, and two perennial and two annual forbs. Twelve of the species had their lowest biomass and lowest survivorship at water depths greater than 0 cm. The root:shoot ratio, however, did not change across water-depth treatments. Biomass accumulation differed by plant form (sedg- esϾforbsϾgrasses). Annuals had the greatest biomass values across the widest range of water depths com- pared to perennials. Of the fourteen plants tested, Lythrum salicaria (purple loosestrife), one of the two invasive, non-native species tested, had the greatest biomass at water depths from Ϫ6toϩ2, whereas Phalaris arundinacea (reed canarygrass), the other invasive, had comparatively small mean biomass values. Ranking of biomass between species was highly concordant between non-flooded treatments but not significantly concordant between flooded treatments indicating that plant species have distinct responses to flooding. This research suggests that newly established plant seedlings in wetland restorations should not be submerged, or if submergence is unavoidable, annuals and sedges may be more tolerant of prolonged flooding. Key Words: wetland restoration, flooding, hydrology, fundamental niche, hydrophyte INTRODUCTION Changes in land use have resulted in destruction and fragmentation of wetlands in North America over the Much of the literature in wetland research is focused past century, which has affected water quality, water on how hydrologic regimes affect the structure and recharge, and wildlife habitat. As a result, wetland res- function of a given wetland (Spence 1982, Gerritsen toration has recently become a common and wide- and Greening 1989, van der Valk et al. 1994, Baren- spread activity. Wetland restoration research has fo- dregt et al. 1995, Owen 1995, Busch et al. 1998, cused primarily on seed viability, seed germination, Moore et al 1999, Keddy and Fraser 2000, Baldwin et and seed-bank dynamics, as they relate to hydrology, al. 2001, Johnson et al. 2004). Laboratory, field, and nutrients, temperature, and time (van der Valk 1981, modeling techniques have all been employed to gain Keddy and Ellis 1985, Schneider and Sharitz 1986, insight into the processes that determine how abiotic Gerritsen and Greening 1989, Budelsky and Galato- factors contribute to wetland plant community struc- witsch 1999, van der Valk et al. 1999, Leck 2003, ture (van der Valk et al. 1994, Ellison and Bedford Jensen 2004). Little information is available regarding 1995, Weiher and Keddy 1995, Casanova and Brock how species perform, once germinated, over a range 2000, Cole and Brooks 2000, Baldwin et al. 2001, De of hydrologic treatments (Weiher et al. 1996, Budelsky Steven and Toner 2004, Johnson et al. 2004). Of all and Galatowitch 2000). Experiments have determined potential abiotic factors, hydrologic regimes have gen- that an increase in water level generally reduces the erally been shown to correlate most strongly with veg- growth and affects morphological responses of wet- etation type (Keddy 2000, De Steven and Toner 2004). land plants (Coops et al. 1996, Newman et al. 1996, 520 Fraser & Karnezis, WATER DEPTH EFFECTS ON PLANT BIOMASS 521 Table 1. Fourteen wetland plant species that were used in the experiment and grown at seven water depths. Scientific Name and Authoritya Code Common Name Life Form Life Cycle 1Calamagrostis canadensis (Michx.) Beauv. Cc Blue-joint grass Grass Perennial 1Carex lurida Wahlenb. Cl Hop sedge Sedge Perennial 1Carex stipata Hudson Cs Sawbeak sedge Sedge Perennial 2Carex tribuloides Wahlenb. Ct Bristlebract sedge Sedge Perennial 1Carex vulpinoidea Michx. Cv Fox sedge Sedge Perennial 2Elymus virginicus L. Ev Virginia wildrye Grass Perennial 2Glyceria canadensis (Michx.) Trin. Gc Rattlesnake mannagrass Grass Perennial 1Lythrum salicaria L. Ls Purple loosestrife Forb Facultative annual 1Mimulus ringens L. Mr Monkey flower Forb Facultative annual 1Phalaris arundinacea L. Pa Reed canarygrass Grass Perennial 1Rumex orbiculatus A. Gray Ro Great water dock Forb Perennial 1Scirpus cyperinus (L.) Kunth. Sc Woolgrass Sedge Perennial 2Verbesina alternifolia (L.) Britt. Va Yellow ironweed Forb Perennial 2Zizania aquatica L. Za Annual wildrice Grass Annual a from Crow and Hellquist (2000) 1 indicates field-collected seed. 2 indicates seed that was purchased from Ernst Conservation Seeds. Lentz and Dunson 1998, Lenssen et al. 1999, Miller short time period. Fourteen wetland species, repre- and Zedler 2003); however, the number of species test- senting a range of growth forms, were grown at seven ed is generally less than five for each study, the water- static water depths, from Ϫ6cmtoϩ6cmin2cm depth treatments tend to differ by at least 5 cm, and increments relative to the soil surface. Two-week-old the plants tested are generally not seedlings but veg- seedlings were used because it is already well-estab- etative ramets of mature plants (but see Kercher and lished that flooding is a constraint to germination (van Zedler 2004). der Valk 1981, Keddy and Ellis 1985, Casanova and Wetland restorations generally involve the new es- Brock 2000, Jensen 2004), but the gap in the literature tablishment of a plant community, either by natural regards the survivorship and growth of seedlings to seed dispersal, introduced seeds, or planted seedlings. water depth. The following questions were posited. (1) The establishment phase of the plant community is Does water depth affect biomass and root:shoot ratios critical to colonization and stand development. Con- of these wetland plants? (2) Is there a difference in trolled mesocosm experiments have shown that control seedling survivorship between water depths and be- of water depth during the first year can significantly tween species? (3) Does biomass at different water influence the potential for growth, mortality, and stand depths differ between life forms (grass, forb, and development in later years (Weiher and Keddy 1995, sedge) and life cycles (perennial and annual)? (4) Does Budelsky and Galatowitsch 2000). the hierarchy of growth performance remain consistent In natural systems, isolating an individual environ- across water-depth treatments? (5) Do non-native in- mental factor’s effect upon that system is nearly im- vasive species (Lythrum salicaria and Phalaris arun- possible. Using laboratory techniques, we can more dinacea) produce more biomass across the seven water clearly delineate the role of hydrology on a number of depths compared to the twelve native species tested? species (Fraser and Keddy 1997). Isolating the effect of water depth on a range of wetland species can re- MATERIALS AND METHODS veal fundamental niche boundaries, information that can be applied to predictive models of plant establish- Twelve species were selected to represent a broad ment and wetland restoration. range of plant life forms typically found in temperate The purpose of this experiment was to investigate eastern North American marsh and wet meadow wet- the growth responses (measured as seedling survival lands (Crow and Hellquist 2000) that are commonly and accumulated biomass) under different water used in wetland restoration projects (Hammer 1997, depths at small (2 cm) increments in controlled micro- Cronk and Fennessy 2001) (Table 1). In addition, we cosms. Although static water levels are generally not included two invasives, Lythrum salicaria (purple normal or realistic conditions found in natural marshes loosestrife, a facultative annual forb) and Phalaris or wet meadows, we were interested in identifying the arundinacea (reed canary grass, a perennial grass), constraints of absolute water depth to initial establish- commonly found throughout North America (Apfel- ment and performance of seedlings over a relatively baum and Sams 1987, Thompson et al. 1987). Species 522 WETLANDS, Volume 25, No. 3, 2005 represent three life forms (forbs, sedges, and grasses) species relating to germination date, but no more than and two life cycles (facultative or obligate annuals and two months separated the first and last species. perennials). Seeds were either purchased through Ernst Seed Company (Meadville, PA) or collected in the Standardized Conditions field. The seed source from Ernst Seed Company is a combination of field-collected and cultivated plants In order to isolate the effects of the water-depth that are bulk processed so that there is genetic vari- treatment on the fourteen species better, other abiotic ability. Field-collected seed were taken from multiple factors were standardized. A daily, 14-hour photope- individuals and from at least three sites, which were riod was provided