J. Aquat. Manage. 46: 117-119 NOTES Effects of Water Level Fluctuation on americana Michx Growth

CHETTA S. OWENS1, R. M. SMART1 AND G. O. DICK1

INTRODUCTION 122 cm. Water level was maintained by continuous low-flow from nearby Lewisville Lake. Wild ( Michx.) is a native, sub- In July 2003, six containers were harvested from each mersed found in the eastern depth, processed into aboveground and belowground mate- westward to and south throughout the Gulf rial, and dried at 55 C in a Blue M forced-air oven (General Coast states (Korschgen and Green 1988, USDA 2007). An Signal, Atlanta, GA) to a constant weight. Rosettes (daughter important food source for waterfowl and aquatic mammals ) were counted and maximum length was mea- (Fassett 1957), wild celery also provides habitat for fish, sedi- sured. At this time, remaining containers from the four ment stabilization benefits, improved water quality and clari- depth treatments were placed at a same depth (91 cm) to ty, and can slow or prevent the invasion of nonindigenous evaluate effects of increasing, stable, and decreasing water aquatic plant species (Smart 1993, 1995, Smart et al. 1994, levels on wild celery growth. After four weeks (August 2003), Smart and Dick 1999). containers were harvested and processed as described above. Numerous constructed reservoirs in the United States are Rosette number, maximum length, aboveground, and be- devoid of aquatic vegetation, exhibit some degree of water lowground biomass means were compared using a one-way level fluctuations, and lack propagule sources for timely nat- ANOVA and LSD (least significant differences) test at p = ural establishment of wild celery to occur (Smart 1993, 1995, 0.05 level of significance (Statistix Analytical Software, Talla- Smart et al. 1994, Smart and Dick 1999). Consequently, wild hassee, FL). Analyses were performed on data from both July celery plants are widely used in efforts to introduce native and August harvests and between each depth change. plants into manmade reservoirs as well as to restore natural lakes that have been negatively impacted by human activities RESULTS AND DISCUSSION or environmental conditions. Because of the importance of wild celery in these restoration activities and the likelihood Significant differences occurred between the shallowest that systems in which it is planted will experience water level depth (18 cm) and all other depths for both rosette num- fluctuations, a basic understanding of wild celery growth re- bers and maximum leaf lengths (Figure 1) at the July har- sponse to varying depths is crucial. vest. Rosette numbers decreased by more than 50% as depth The objectives of this study were to (1) determine wild cel- increased, indicating wild celery was not allocating resourc- ery rosette (daughter plant) number and biomass produc- es to localized expansion at greater depths. Smart et al. tion at four depths, and (2) determine changes in rosette (1994) found that wild celery was capable of adjusting bio- number and biomass production when exposed to fluctuat- mass allocation and morphology to maximize photosynthe- ing depths. sis, and that under lower light conditions fewer rosettes were produced. Maximum leaf length increased with depth METHODS increases, showing a 5-fold increase between 18 cm and 122 cm by July, likely a response to lower light conditions at Six wild celery plants (average 5 per plant) were greater depths (Titus and Stephens 1983, Sculthorpe 1985). planted into each of 40 6.7-L containers (diameter 35.6 cm, Aboveground biomasses for the July harvest were significant- depth 10.2 cm) of sterilized pond sediment amended with ly different between the most shallow depth (18 cm) and 1.4 g ammonium sulfate in December 2002; sediments were plants grown at the two deepest depths (91 and 122 cm). collected from a pond at the Lewisville Aquatic Ecosystem Aboveground biomass production, grown at different Research Facility, Lewisville, Texas (LAERF). Ten containers depths resulted in plants allocating available resources dif- per depth were randomly distributed atop a concrete pad on ferently via local expansion or elongation. Belowground the bottom of a LAERF pond at four depths: 18, 46, 91, and biomasses were not significantly different among depths in July. -to-shoot ratio indicated that plants growing at the deeper depths sacrificed belowground production () 1U.S. Army Corps of Engineers Engineer Research and Development to increase aboveground biomass production (Figure 2). Center-Lewisville Aquatic Ecosystem Research Facility, 201 Jones St., Lewis- ville, TX 75056. Received for publication September 26, 2007 and in revised After all wild celery plants were transferred to the same form December 12, 2007. depth (91 cm), the root-to-shoot ratio stabilized around 0.5,

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Figure 2. Effects of depth change on root-to-shoot ratio in wild celery.

fold to reach the water surface and widened (Owens, pers. observ.), becoming more similar to those grown in deeper wa- ter. Although leaf length remained significantly shorter than that of the greatest depth (122 cm), this difference was likely attributable to leaves produced by the deeper plants before being moved to 91 cm depths. Rosette numbers increased (30%) in plants moved from 122 cm to 91 cm depths. After the transfer to the same depth (91 cm), biomasses of all treat- ments became similar in August, although significant differ- ences were still found between plants grown at the previously more shallow depths (18 and 46 cm) when compared to Figure 1. Productivity of wild celery at different depths and the effect of plants grown at deeper depths (122 cm). Leaf size, shape, and change in depth (from original depth to 91 cm) on productivity. Graph bars represent the mean of 6 or 4 containers (in July and August, respectively) area are strongly influenced by depth and shading (Spence et with standard error bars. Lower case letter indicates significant differences al. 1973, Titus and Stephens 1983, Madsen 1991). between July sampling. Number indicates significant differences between Although this study did not evaluate light availability at August sampling, and * indicates significant differences between July and each depth, visibility was generally clear to the bottom of the August sampling for depth change. (A) Average rosette number for July (p = pond. Effects of depth on resource allocation and resultant 0.0000), August (p = 0.0118) and depth change to 91 cm from 18 (p = 0.0022), 46 (p = 0.0102), 91 (p = 0.0196) and 122 (p = 0.0144) cm. (B) Max- change in growth patterns were likely due to differing light imum leaf length (cm) for July (p = 0.0000), August (p = 0.0347) and depth availability at each depth. Significant differences were detect- change to 91 cm from 18 (p = 0.0000), 46 (p = 0.0004), 91 (p = 0.0048) and ed for aboveground biomass in August (Figure 1), although 122 (p = 0.2703) cm. (C) Aboveground dry weight (g) for July (p = 0.0643), no differences were detected for belowground biomass. August (p = 0.0460) and depth change to 91 cm from 18 (p = 0.0316), 46 (p = 0.2952), 91 (p = 0.3843) and 122 (p = 0.0030) cm. (D) Belowground dry Although water levels may impact establishment and weight (g) of wild celery for July (p = 0.0582), August (p = 0.4093) and growth of wild celery, this study indicates that plants can re- depth change to 91 cm from 18 (p = 0.0557), 46 (p = 0.0311), 91 (p = spond to changes in depth (or light penetration) by potential- 0.0007), and 122 (p = 0.0000) cm. ly altering rosette production (122 cm depth) or leaf length (18 and 46 cm). While we had only one treatment (122 cm) that decreased in water level, Titus and Adams (1979) found except for the plants that had been growing at the 122 cm that wild celery could respond to light change within 24 h, depth (Figure 2). supporting the findings in this study that found wild celery ac- Statistical differences occurred between months (July and climating rapidly to changing light. This information might be August for same treatment, different depth) for most parame- useful in ascertaining best planting depths for wild celery colo- ters (Figure 1), and these differences were driven primarily by nies during restoration projects, especially where water level changes in depth, elongation over time (even at the same fluctuations are predictable. The ability of wild celery to toler- depth), and plant maturation. Four weeks after wild celery ate water level fluctuations and different water depths indi- grown at different depths were moved to the same depth cates the importance of wild celery for restoration efforts. (91 cm) plants shifted resource allocation and exhibited changes in growth patterns (Figure 1). Rosette numbers in ACKNOWLEDGMENTS the shallowest depth plants (18 cm) decreased by almost 40% after greater inundation due to self-thinning. As self-thinning This research was conducted under the U.S. Army Corps occurred, rosette numbers decreased, leaves elongated by 4- of Engineers Aquatic Plant Control Research Program, U.S.

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Army Corps of Engineers Research and Development Center Smart, R. M., J. W. Barko and D. G. McFarland. 1994. Competition between under the program leadership of Robert Gunkel. Permission verticillata and Vallisneria americana under different environmen- tal conditions. Technical Report A-94-1, U.S. Army Engineer Waterways to publish this information was granted by the Chief of Engi- Experiment Station, Vicksburg, MS. 34 pp. neers. We would like to thank LeeAnn Glomski and Joe Snow Smart, R. M. 1995. Preemption: An important determinant of competitive for review of this paper. success. Miscellaneous paper A-95-3, U.S. Army Engineer Waterways Experiment Station, Vicksburg, MS. pp. 231-236. Smart, R. M. and G. O. Dick. 1999. Propagation and establishment of LITERATURE CITED aquatic plants: A handbook for ecosystem restoration projects. Technical Report A-99-4, U.S. Army Engineer Waterways Experiment Station, Fassett, N. C. 1957. A manual of aquatic plants. University of Wisconsin Vicksburg, MS. 34 pp. Press, Madison. 405 pp. Spence, D. H. N., R. M. Campbell and J. Crystal. 1973. Specific leaf area and Korschgen, C. E. and W. L. Green. 1988. American wild celery (Vallisneria zonation of freshwater macrophytes. J. Ecol. 61:317-328. americana): Ecological considerations for restoration. U.S. Fish and Wild- Titus, J. E. and M. S. Adams. 1979. Coexistence and the comparative light life Service, Fish Wild. Tech. Rep 19. 24 pp. relations of the submersed macrophytes Myriophyllum spicatum L. and Val- Madsen, J. D. 1991. Resource allocation at the individual plant level. Aquat. lisneria americana Michx. Oceologia 40:273-286. Bot. 41:67-86. Titus, J. E. and M. D. Stephens. 1983. Neighbor influences and seasonal Sculthorpe, C. D. 1985. The biology of aquatic vascular plants. Edward growth patterns for Vallisneria americana in a mesotrophic lake. Oceolo- Arnold Ltd., London, England. 334 pp. gia 56:23-29. Smart, R. M. 1993. Competition among introduced and native species. Proc. USDA, NRCS 2007. The PLANTS Database (http://plants.usda.gov, 2007). 27th Annual Meeting, Aquatic Plant Control Research Program. Miscella- National Plant Data Center, Baton Rouge, LA. neous paper A-93-2, U.S. Army Environmental Research and Develop- ment Laboratory, Vicksburg, MS. pp. 235-241.

J. Aquat. Plant Manage. 46: 119-121 Effect of Water Temperature on 2,4-D Ester and Carfentrazone-ethyl Applications for Control of Variable-leaf Milfoil

LEEANN M. GLOMSKI1 AND MICHAEL D. NETHERLAND2

INTRODUCTION Two herbicides that have been shown to effectively control variable-leaf milfoil include 2,4-D ester ([2,4-dichlorophe- Variable-leaf milfoil (Myriophyllum heterophyllum Michx.) is noxy]acetic acid) and carfentrazone-ethyl (a,2-dichloro-5-[4- a submersed plant native to southwestern Quebec and On- (difluoromethyl)-4,5-dihydro-3-methyl-5-oxo-1H-1,2,4-triazol- tario, to North Dakota and southward to New and 1-yl]-4-fluorobenzenepropanoic acid, ethyl ester). In green- Florida (Godfrey and Wooten 1981). In the Northeastern house studies, 2,4-D ester at 500 and 1500 µg ai L-1 exposed for U.S., variable-leaf milfoil is not native and is considered an 3, 8, and 24 hours provided 98 to 100% control of variable-leaf invasive and weedy species. As an invasive species, it causes milfoil (Netherland and Glomski 2007). Bugbee et al. (2003) many of the same problems as Eurasian watermilfoil (Myrio- also reported that 227 kg ha-1 2,4-D ester as Navigate con- phyllum spicatum L.), including shading out native submersed trolled nearly all the variable-leaf milfoil in treated field sites. vegetation and interfering with recreational activities and wa- Carfentrazone at 100 µg ai L-1 for 6 to 30 hours was reported to ter supplies (Halstead et al. 2003, NH-DES 2002). Variable- provide 61 to 81% control of variable-leaf milfoil. Doubling leaf milfoil is an aggressive invader that can grow up to one the rate of carfentrazone did not improve efficacy (Glomski inch per day under optimal nutrient, temperature, and light and Netherland 2007). While there is no published literature conditions and spreads mainly via fragmentation (NH-DES regarding field applications of carfentrazone to control vari- 2002). able-leaf milfoil, recent field trials in North Carolina have demonstrated good control (Rob Richardson, pers. comm.). The effect of water temperature on efficacy of aquatic her- 1U.S. Army Engineer Research and Development Center, Environmental bicide applications is not well documented in the literature. Laboratory, 201 E. Jones St., Lewisville, TX 75057; e-mail: LeeAnn.M.Glom- Westerdahl and Getsinger (1988) suggest that aquatic plants [email protected]. have low metabolic activity in cooler waters, and this can in- 2 U.S. Army Engineer Research and Development Center, Center for hibit herbicide uptake. Studies done by Netherland et al. Aquatic and Invasive Plants, Gainesville, FL 32653. Received for publication July 22, 2007 and in revised form December 12, 2007. (2000) and Poovey et al. (2002) demonstrated that as water

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