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A . . ‘ ‘ 4" 8.095 (35/4443 This is to certify that the thesis entitled EFFECTS OF SALINITY ON THE BIOTA OF NATURAL AND CREATED WETLAND COMMUNITIES presented by CYNTHIA KOPPEN HODGES has been accepted towards fulfillment of the requirements for the MS. degree in ZOOLOGY 27¢ moi/4’: Major Professor’s Signature MW / Date MSU is an Affirmative Action/Equal Opportunity Institution LIBRARIES MICHIGAN STATE UNIVERSITY EAST LANSING, MICH 48824—1048 fl,._.._.-.-—.-.—._--.-o-n-o-~ — ..__ a. —« r V —-—~'- r—fi PLACE IN RETURN Box to remove this ched50°/o frequency of occurrence and relative abundance and 15 common plants meaning those with >50% frequency of occurrence but <50% relative abundance) were analyzed individually, and all other plants were grouped into an “other species” category. Averages and standard deviations for environmental variables were determined and differences in these variables based on the presence or absence of S. americanus were determined using a t-test in Systat 8.0. Abundances of each plant species were correlated to the environmental variables and to abundances for each year using a Spearman’s rank correlation in Systat 8.0, since the data were not normally distributed (Wheater and Cook 2000). To determine environmental variables that could be used in a regression analysis, a principal components factor analysis was run using Systat 8.0 to remove the collinear data in the set of environmental variables (Kachigan 1986). Environmental factors that were chosen based on factor analysis were standardized and used in forward stepwise multiple regression analyses for dominant, common and other plant species for each year using Systat 8.0, although the environmental factors were only recorded in 2001. To determine changes in plant community over time, a Wilcoxon signed rank test of the differences between years was used to compare the stem counts/m2 and relative abundances of species per plot for 2000 and 2001. The Wilcoxon signed rank test was used because of expected non-normal distribution of the data (Schabenberger 1999). Significant differences were determined at p<0.05. Btu!!! The species found at the Maple River salt seep are listed in Appendix B. The dominant species were Schoemplectus americanus and Aster Ianceolatus in the saline area and T ypha spp. around the edges and to the north of the saline area (Figure 1a). 16 Mentha arvensis was also considered common; however, its relative abundance was low (Figures 2 and 3, Table 1) in most plots. Two species of endangered, native Michigan halophytes were found at this seep (MNFI, Catling and McCay 1981): S. americanus was abundant both in 2000 and 2001, while Eleocharisparvula was found only around the seep area in 2001. In 2000, a few dead Chara spp. plants, the third native Michigan halophyte, were found at the seep area, but none were found in 2001. A total of 41 species were found in the plant sampling. Fifteen species were found in the S. americanus patch, but six of these species were found only on the edges of the patch. Fifieen species were also exclusively found in the upland forest, six species were exclusively found in the forested wetland and one species was exclusively found in the T ypha patch (see App. B). The means for all plots of the stem counts/m2 and relative abundances for the dominant and common plant species were the same for all species except S. americanus and 71 latrfolia (Table 1). Both species of T who were grouped together in 2000, but were separated into two species in 2001. 71 angustrfolia was significantly correlated with the grouped ijha spp. for both years since it was found more often than T. latifolia in 2001. Correlations comparing 2000 to 2001 showed positive r, meaning similar patterns of association for all species except I Iatrfolia in 2001 to T who spp. in 2000 (Table 1). Relative abundance per plot (Figures 2 and 3) were significantly correlated to average stem counts per m2 for each dominant plant species within the same year (Table 2). Correlations for stem counts (Figures 4-8, Table 2) and relative abundance (Figures 2 and 3) comparing 2000 to 2001 were also significant for all the dominant species except 17 T. larifolia. The number of species per plot (Figure 9) positively correlated with stem counts of other species (Table 2). Conductivity data (Figure 10), chemical data for C1, Ca, and Na (Figures 11 and 12) and depth to water table measures (Figure 15) confirm that the open muddy area assumed to be the primary salt seep (0 point in the graphs) is the source of most salinity in this wetland, although there seems to be a secondary seepage area at a distance of about 50-55 m fiom point 0 along transect A (Figures 11, 12 and 15). There also is a peak in nearly all water chemistry measurements (except N03-N, Figure 13) at —40m on transect A (Figures 11-14), however, this peak is not seen in the conductivities (Figure 10) that were performed in the field. The peak stem counts and relative abundances for S. americanus were near the open muddy seep area, and 60 to 80m fiom the muddy area on transect A (Figures 2-4). These high counts of S. americanus corresponded to the peak conductivities, Cl, Ca, and Na concentrations (Figures 10-12). The depth of the water table, conductivity, alkalinity, Ca, Na, and C1 were significantly higher in plots where S. americanus was present (Table 3). However, temperature and N03 were significantly higher in plots where S. americanus was absent (Table 3). Spring water depth, pH, N114, ortho-P, K, and Mn were similar whether S. amerims was present or absent (Table 3). Conductivity was measured in both spring and summer of 2001. The trends in conductivity for spring and summer 2001 were similar for both transects (Figure 10). Conductivities were generally lower in the spring than in the summer, perhaps due to a higher water table and/or the influence of spring flooding. In particular, the northern 18 section of transect B was strongly influenced by the flooded Maple River in the spring, and conductivities for this part of the transect were much lower tlnn those recorded in the fall (Figure 10). Environmental factors indicated strong positive correlations between Na, C1, Ca and alkalinity (Table 4, Figure 17). Spring water depth, N03, and ortho-P (Figures 13, 14) were also positively correlated to each other, but negatively correlated to Na, Cl, Ca and alkalinity (Table 4, Figure 17). Spring water depth was also negatively correlated to conductivity and depth to water table, but conductivity and depth to water table were positively correlated to each other (Table 4, Figure 17). Temperature was negatively correlated to conductivity, and ortho-P was positively correhted to NIL. K, Mn, and pH had no significant correlations (Table 4, Figure 17). Table 5 shows the correlations of the plant data to the environmental data. S. amerr'canus was positively correlated with depth to water table, conductivity and Na, but negatively correlated with spring water depth, N03, and ortho-P in both years for stem counts/m2 and relative abundance per plot (Table 5). M. arvensis stem counts/m2 were positively correlated with alkalinity (Table 5). Other species were negatively correlated with depth to water table for stem counts/m2 and for 2000 relative abundance per plot (Table 5). Based on the principal components factor amlysis, four environmental factors were determined to account for the majority of variation. These factors were depth to water table, conductivity, pH, and alkalinity. However, since I was interested in effects of salinity, I also included C1 in the analysis. Also, nutrients are important factors in 19 plant communities, so N03 and ortho-P were also included. Thus, a total of seven environmental factors were used in the regression analysis (Table 6). Stem counts and relative abundance of S. americanus were positively related to conductivity and negatively related to ortho-P for both years. S. amerr'canus was also negatively related to Cl, N03, and alkalinity in 2000, but not 2001. In 2000, S. americanus stem counts were negatively related to depth of the water table, but in 2001, stem counts and relative abundance were positively correlated to depth of water table (Table 6). M. arvensis was negatively related to alkalinity for all but 2001 relative abundance, and positively related to C l in 2000, but not in 2001 (Table 6). A. Ianceolatus relative abundance was positively correlated to ortho-P for both years (Table 6). In 2001, T. Iatifolia was positively correlated to ortho-P for stem counts and relative abundance. T. angustrfolia was negatively correlated to conductivity in 2001 (Table 6). Other species were negatively related to alkalinity in all but relative abundance for 2000. Other species were also negatively related to conductivity in 2000, but not 2001, and negatively related to depth to water table in 2001, but not 2000 (Table 6). D ion The water levels at the muddy open area were never deeper than a few centimeters. Even when the area was not flooded, the water table was always within a few ems of the surface, and the soil remained saturated for the whole growing season. Chemical and environmental data confirmed that this muddy area was the primary source of saline water into the wetland, although a secondary seepage area appeared to be likely about 50 to 60 m to the southeast along transect A (Figure 1b) based on chemical data (Figmes 10-12 and 15). The wetland around the primary seep maintained a fairly stable 20 water level throughout the year ranging fiom 5 cm above to 17 cm below the soil surface. The areas down slope to the north, east, and west, where the plant community visibly changed fiom a S. americanus dominated community to a T wha dominated or forested wetland community, had much greater fluctuations in water depth, ranging from nearly one m deep in the spring to 30 cm below the soil surface in late summer. The areas down slope of the S. americanus patch were mainly flood plain wetlands, and, not surprisingly, water level appeared to be controlled by the flood regime of the Maple River. The seep was more dependent on ground water, as was indicated by its more consistent water levels. This seepage area is near the base of a steep hill or small escarpment that separates the adjacent upland from the floodplain. The upland area south of the seep never had water near the soil surface. Of the 41 species found in the seep plant sampling, only 15 were found in the S. amerr’camrs patch (App. B). Of these 15 species, 6 were at the edges of the patch and 9 were found within the patch. Nine species found were salt tolerant, and eight of the salt tolerant species were found in the S. americanus patch (Peltandra virginica, an introduced species in Michigan, was only found in the forested wetland, App A and B). Two of the species found in the patch are rare and endangered native halophytes of Michigan (S. americanus and E parvula, Catling and McKay 1981, MNFI). Atriplex panda, an introduced, non-native salt tolerant species in Michigan (Voss 1985), was frequently encountered within the S. americanus patch, and II mrgustrfolr’a, another introduced halophyte (Chapman et al 2001), was abundant on the edges and formed monodominant stands on the north end of the S. americamrs patch (App B). The four remaining salt tolerant species (Aster lanceolatus, T. latrfolia, Mentha arvensis, and 21 Acorns calanms) are common throughout the state of Michigan (Voss 1972, 1996). Although the remaining seven species are not identified as salt tolerant, four of these species (Eupatorium perfoliatum, E. maculatum, Onoclea sensitiva, and Equisetum spp.) were found in another salt seep in Michigan (Chapman et al 1985) and are often found along roadsides (V oss 1996, Billington 1952) indicating possible salt tolerance. These remaining seven species (Scuttelarr’a galericulata, Phalaris amndinacea and Pilea spp. in addition to the four mentioned above) found in the S. amerr'aamrs patch (App. B) are widespread and common throughout the state of Michigan with the exception of PiIea spp. which is found mostly in southern Michigan. In most cases, the plants that were found at the seep are common species found throughout Michigan, but were found in low numbers in the seep (except for the endangered halophyte E parvula). The four dominant and common species at the seep were all salt tolerant, and with the exception of S. amerr‘ccnms are also very common in the state of Michigan. Based on my literature search, 88 native plant species in Michigan and numerous introduced species have salt tolerant populations at some point in their habitat range (Chapman et al 2001, Winchell 1860, Voss 1972, 1985, and 1996, Reznicek 1980, App. A). However, only nine salt tolerant species were found at the seep, two introduced and the other seven native Michigan species (App A and B). Furthermore, two endangered Michigan halophytes (MNF I) were found, one of them dominating a large portion of the seep. Wheeler (1891) reported these two species at this seep (a “deer lick” near Hubbardston) in the late 18003. Thus, this seep appears to be a well-suited habitat for these two rare species in Michigan. 22 Abundance of S. amerr‘canus was related to conductivity. S amerr‘ccmus is a halophytic plant, and as expected was associated with high ion concentrations. E. parvula, another native Michigan halophyte, was very rare and found only in the open, non-vegetated seep area in 2001. E. parvula seedlings emerge in greater densities from non-vegetated areas (Baldwin et al. 1996) as was supported by this study. Although this species is capable of germinating at high salinities (up to 16 ppt in flooded conditions), salinities of only 2 ppt can greatly reduce germination (Baldwin et al. 1996). Although this wetland is predominately ground water fed, rain can reduce the salinity of surface layers by 1% salinity (Chapman 1974). The annual precipitation (based on monthly precipitation data) from Grand Rapids, MI (July 1997-current) for both years of this study were nearly the same, but a few years before 2000, the precipitation was much lower. Perhaps, salinity in this low precipitation period rose to levels high enough to eliminate Epw'vulafiomthe seepagearea Ifso, it is possiblethatEparvuIa seeds may have required an extended period of two or more years of average or higher than average rainfall to lower salinities in the seep area to levels low enough for germination. This is a possible explanation of why germination was only detected in 2001 and not in 2000 despite similar level of rainfall in both years. Since I did not collect chemical data in 2000, my data were not sufficient to document that salinities were indeed lower in 2001 than in 2000. Since E pcn'vula is such a rare plant in Michigan (MNFI, Voss 1972), and the seep I studied is only one of two areas in Michigan where both threatened species (S. mnericanus and E parvula) of native halophytes grow, firrther studies should be conducted on its life cycle and habitat needs in order to develop a management plan to preserve the species in Michigan. 23 S. americamrs abundance was also negatively conelated with ortho-P. This species is a sub-climax species (Broome et al. 1995), meaning it is easily out-competed by other species. In 2001, T latifolia abundance was correlated to ortho-P; however, Twin spp. grouped together in 2000 and Ti mgustrfolia in 2001 did not have significant conelations with ortho-P. Since the species of T win were not distinguished in 2000, correlations to environmental factors may have been hindered. T win spp. (especially T. Iatrfolia) are known to be highly competitive, and will take over areas that are enriched with nutrients (Svengsouk and Mitsch 2001, Hutchinson 1975). T laafolia was not found in this wetland in a 1983 survey (Shaddalee 1983), and the T win dominated stand where this species is predominately found was also not indicated in the 1983 study (Shaddalee 1983). T latrfolia is a very competitive species that often displaces T. wtgus'tifoha to less desirable deep water (Grace and Wetzel 1981) and high salinity habitats (McMillan 1959). Maintaining this S. americanus population may depend on maintaining low P levels in and down slope of the saline water seepage area. The composition of the plant community did not appear to change fi'om 2000 to 2001 except that E parwla was present in 2001 but not in 2000. In comparing field notes of this wetland fi'om 1983 (Shaddellee 1983) with my study, the plant community structure appears to be similar, especially in terms of the distribution of the dominant species present. S mnericanus was the dominant plant in 1983, with 71 mtgusafolia being locally dominant on the north, east and the eastern portion of the south edges. However, I found a mixed patch of both T. angustrfolia and T. Ianfolia on the south west edge of the S. americcmus dominated patch that was not indicated in Shaddellee’s notes (1983). In both studies, Aster lanceolams (called A. simplex, see Voss 1996) was 24 abundant to the point of being considered co-dominant. In both surveys Eupatorium macularum, E. perfoliatum, M. m'vensis, Onoclea sensibilis, Atriplex hastata, Tirebpteris palusmls and Pilea spp. were present in lower abundances. A few of the rarer species in both surveys did not correspond (Shaddellee 1983). However, being rare, those species could have escaped notice or recently immigrated into or emigrated out of the wetland. Also, rare species are likely to differ between any two studies of the same area when random quadrat placement within plots is part of the experimental design especially when limited numbers of quadrats are sampled per plot (e. g. 3 in this study). The greatest difference between the two surveys was that Shaddellee (1983) found Eleocinris rostellata to be co-dorninant with S. americamrs in this wetland, and that species was not found in my study 17 years later. Wheeler (1891) also found this species at the seep over 100 years ago. E rostellata is an early colonizer of marl beds and seems to grow best on marl soil that receives runoff fiom glacial till (Seischab et al. 1985). Since this wetland is groundwater fed, E rosrellata may have been less hearty and unable to compete with the abundant and dense population of S. americanus. However, since E. rostellata was likely present in this seep from Wheeler’s (189]) observations to Shaddellee (1983) study, its absence would indicate a recent disappearance. Since Shaddellee (1983) reported E rostellata as a codominant species with S. americwrus it is unlikely that my survey would have missed this species even though I did not sample the entire wetland. This species absence may be an early indicator of a changing environment at the seep, and the seep should be continually monitored for changes in the plant assemblage. 25 Cl, Ca, Na, and K were the four highest concentrated ions, respectively, at this seep. Cl, Na, and K ion concentrations were similar to those found in seawater (Fortescue 1980). Unfortunately, Mg another common salt of seawater was not tested in this study. Based on the geology of Michigan, this wetland is very likely obtaining water fi'om the saline aquifers that are remnants of the Paleozoic seas that covered Michigan (Don and Eschman 1970). Ca, which tends to be in low concentrations in sea water (F ortescue 1980) and in the brine waters of Michigan (Cook 1914, Dorr and Eschman 1970, Winchell 1860), likely derives from the glacial till deposits in the lower peninsula which tend to have high concentrations of calcium carbonates (Chapman et al. 1985). Compared to the water chemistry of bogs, swamps and fens in Northern Michigan studied by Schwintzer and Tomberlin (1982), C1, Ca, Na, K, conductivity, and pH were all higher at the Maple River salt seep. The alkalinity and ortho-P (reactive-P) at the Maple River salt seep wetland were comparable to those found by Schwintzer and Tomberlin (1982) for northern Michigan wetlands. However, the N03 and NH4 concentrations at the salt seep were more similar to river water (Allen 1995) than to Northern Michigan wetlands (Schwintzer and Tomberlin 1982), since the N03 concentration were much higher than the NH4 concentrations at the Maple River salt seep. The Michigan Department of Environmental Quality (MDEQ 2003) studied the river water of the Maple River watershed. They found that the river water had ammonia values ranging fi'om 0.012 to 9.71 mgN/L and N03 levels ranged fiom 0.09 to 14.1 mgN/L (MDEQ 2003). The NH4 and N03 concentrations at the salt seep were within the range of concentrations found in the river water. Since three edges of the salt seep 26 were flooded by river water, the river water likely contributes its nutrient concentrations to the salt seep wetland. Compared to the salt marsh studied by Chapman et al., (1985), my seep had weaker brine, however, I determined ion concentrations fi'om the water, whereas Chapman et al. (1985) used dry soil samples. Calcium, magnesium and chloride were the most abundant ions, respectively, at the Clmpman et al. (1985) marsh, where as chloride, calcium, and sodium had the highest concentrations, respectively, at my seep. Also pH had a greater range in their study (Chapman et al. 1985), than in my salt seep. This unique and rare salt seep wetland appears to support a healthy population of rare and endangered halophytes in Michigan (MNFI). However, compared to a survey from 17 years ago (Shaddellee 1983), some changes in plant assemblages have occurred. This seep should continue to be monitored and protected to maintain its integrity and to preserve the diversity of the unique plants it contains. 27 Works Cited Allen, J. D. 1995. Stream Ecology. Chapman and Hall, London. Baldwin, A. H., K. L. McKee, and 1. A. Mendelssohn. 1996. The influence of vegetation, salinity, and inundation on seed banks of oligohaline coastal marshes. Am. J. Bot. 83(4): 470—479. Balson, J. 1988. QuickChem Method No. 10-107-04-1-A. Lachat Instruments. Billington, C. 1952. Ferns of Michigan. Cranbrook Institute of Science. Bloomfield Hills, Michigan. Broome, S. W., I. A. Mendelssohn and K. L. McKee. 1995. Relative grth of Spartr'na patens (Ait.) Muhl. and Sa'rpus olneyi gray occurring in a mixed stand as affected by salinity and flooding depth. Wetlands 15: 20-30. Catling, P. M. and S. M. McKay. 1981. A review of the occurrence of halophytes in the eastern Great Lakes Region. Michigan Botanist 20: 167-179. Chapman, K. A., V. L. Dunevitz, and H. T. Kuhn. 1985. Vegetation and chemical analysis of a salt marsh in Clinton County, Michigan. Michigan Botanist 24: 135- 144. Chapman, K. D., L. A. Masters, M. R Penskar, A. A. Reznicek, G. S. Wilhelm, W. W. Brodovich, K. P. Gardiner. 2001. Floristic quality assessment with wetland categories and examples of computer applications for the state of Michigan. 2'“l ed. Michigan Department of Natural Resources. “Wildlife Division. Natural Heritage Program. Chapman, V. J. 1974. Salt Marshes and Salt Deserts of the World. 2”“ edition. Verlag Von J. Cramer, Germany- Cook, W. C. 1914. The brine and salt deposits of Michigan. Mich. Geol. and Biol. Survey Publ 15, Geo. Series 12. 188 pp. Dorr, Jr., J. A. and D. F. Eschman. 1970. Geology of Michigan. University of Michigan Press: Ann Arbor. Fortescue, J. A. C. 1980. Environmental Geochemistry. Srpinger-Verlag, New York. Frank, K., D. Beegle and J. Denning. 1998. Phosphorus. in Recommended Chemical Soil Test Procedures for the North Central Region. North Central Research Publication No. 221. 28 Gelderman, R. H., J. L. Denning and R. J. Goos. 1998. Chlorides. in Recommended Chemical Soil Test Procedures for the North Central Region. North Central Research Publication No. 221 . Grace, J. B. and R G. Wetzel. 1981. Habitat partitioning and competitive displacement in cattails (Twha): experimental field studies. Am Nat. 118: 463-474. Houghton, D. 1838. Report of the state geologist. House documents 24: 96-148 in G. N. Fuller, Geological Reports of Douglass Houghton. Michigan Historical Society, Lansing. 1928. Hutchinson, G. E. 1975. A Treatise on limnology, Vol. 3. Lirnnological botany. Wiley, New York. Kachigan, S. K. 1986. Statistical Analysis. Radius Press, New York. McMillan, C. 1959. Salt tolerance within a T wha population. Am. J. Bot. 46: 521-526. Michigan Department of Environmental Quality (MDEQ), Water Division. 2003. A biological survey of the Maple River watershed and selected tributaries Shiawassee, Clinton, Montcalm, Gratiot, and Ionia Counties, Michigan August 2002. MI/DEQ/WD-03/017. ' Michigan Natural Features Inventory (MNFI). Michigan State University Extension. http://web4. m sue. msu . edu/mnfi/home/cfm. Monthly Precipitation Data Grand Rapids, Michigan (July 1997 through current). http://www.x98ruhg.net/gttp97.html. Nelson, D. W. 1983. Determination of Ammonium in KC] extracts of soils by the salicylate method. Communications in Soil Science and Plant Analysis. 14(11): 105 1-1062. Reznicek, A. A. 1980. Halophytes along a Michigan roadside with comments on the occurrence of halophytes in Michigan. Schabenberger, O. 1999. Statistical Methods for Biologists 1, Course Pack. MSU Printing Services, East Lansing. Schwintzer, C. R. and T. J. Tomberlin. 1982. Chemical and physical characteristics of shallow ground waters in northern Michigan bogs, swamps and fens. Am. J. Bot. 69(8): 1231-1239. Seischab, F. K., J. M. Bernard, K. F iala. 1985. Above and Belowground standing crop partitioning of biomass by Eleocharis rostellata Torr. In the Byron-Bergen Swam, Genessee County, New York. Am. Midland Naturalist. 114(1): 70-76. 29 Shaddellee. August 2, 1983. Field notes, N of the Island, Michigan Natural Features Inventory. Svengsouk, L. J. and W. J. Mitsch. 2001. Dynamics of mixtures of Twin laafoia and Scheonoplectus tabemaemontani in nutrient-enrichment wetland experiments. Am. Midl. Nat. 145: 309-324. Voss, E. G. 1972. Michigan Flora, Part I, Gyrnnosperms and Monocots. Cranbrook Institute of Science, Bloomfield Hills. Voss, E. G. 1985. Michigan Flora, Part II, Dicots (Saurm'aceae-Cornaceae). Cranbrook Institute of Science, Bloomfield Hills. Voss, E. G. 1996. Michigan Flora, Part III, Dicots (Pyrolaceae-Compositae). Cranbrook Institute of Science, Bloomfield Hills. Waisel, Y. 1972. Biology of Halophytes. Academic Press, New York. Warncke, D. and J. R. Brown. 1998. Potassium and other basic cations. in Recommended Chemical Soil Test Procedures for the North Central Region. North Central Research Publication No. 221. Weller, M. W. 1994. Bird-habitat relationships in a Texas estuarine marsh during summer. Wetlands 14: 293-300. Wheater, C. P. and P. A. Cook. 2000. Using Statistics to Understand the Environment. Routledge, London. Winchell, A.,l861. First Biennial Report of the Progress of the Geological Survey of Michigan, Embracing Observations on the Geology, Zoology, and Botany of the Lower Peninsula. 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Environmental Means for all S mnericanus S mnericanus T, p values factors plots (n=18) absent (n=5) present (n=13) Depth to water table -11 i 8 -29 i 1 -8 i 3 -l4.47. (cm) >0.001** Spring water depth 24 i 37 64 i 52 9 i 9 2.35, (cm) 0.077 Conductivity (uS) 1560 :1: 1060 534 i 3 1790 i: 1040 -4.35. 0.001” pH 7.45 i 0.22 7.39 i 0.04 7.47 i 0.24 4.19, 0.253 Alkalinity (mg 248 :l: 82 160 i 1 268 :t 77 -5.00. CaC03/L) >0.001** Temperature (°C) 14.1 i 0.9 14.9 i 0.3 13.9 i 1.0 2.91. 0014" N03 (mg NIL) 3.0i3.6 7.51 i018 1.96ir3.13 6.35, >0.001** NH4 (mg N/L) 0.10 :t 0.07 0.13 :1: 0.02 0.09 i 0.07 1.73, 0.105 Ortho-P (mg/L) 0.04 i 0.05 0.06 :1: 0.01 0.04 i 0.06 1.28. 0.221 K (mg/L) 5.4 :t 5.1 3.3 i 4.5 5.8 i 5.3 -0.85, 0.453 Ca (mg/L) 391 i 298 102 i 2.9 458 i 292 -4.39, 0.001 " Mn (mg/L) 0.18 i 0.29 0.05 i 0.04 0.21 :t 0.31 -1.74, 0.104 Na (mg/L) 203 j: 191 12 i 1 247 i 185 -4.58. 0.001" Cl (mg/L) 661 i 580 74 i 4 797 i 561 -4.64, 0.001" :2222 .32. .2222 22. :22- $2.22. :E222- :2 32 a _ 22 .22. :222 :2. :22- ..§.2-E2- :2222. 22 22 .z _ $2 22 m2 :2. s2. 22. £2. 2 :2. $2 :2 =2 _ .2222. .22 .32 .22 :2 s2. :2. :2. 22. 2.25 2 22 :2. :22. :22. 22. £2,222. 22 n2 6 _ :2- 22. :2. 2.2- 22 :2- a2 22. v. 2 22 Q2 22. 22 22 22- ;2. 2-22 _ :22 :2 :22- 22 2.2. :2- 2.82 _ :2 £22.22- :32- ::2. 2.25% 2 22- a2- .22. $2. 23235. _ 22 22 22 22:2 _ :2- ;2 :2 _ .22 2.82:8 _ 038 523 9 595 22 as 286 22 £2 222 $22 $2 2% ea. 22 2322.2... 32.2 :2 25.2223 225 2.22 2.82 22 .352 2532 £32228 9&2 .56 v a 8:3me 3.. 28 mod v a 8:8me _.. .2822 3552.835 wEEmEoQ 32:26:00 x25 ggoum .v 2an 35 :2- :2- :2- .22 R2. 2 :2 22 $2 22 222-22- 52- :22- 2:25 22- 22- 22 22 e2- 22- 22 22 $2 22 2:2- 22- 22- 22- 22.5 22 22 22 22 $2 22- 22 :2- .2222- .2222 2 2 238.2... 22 :2 R2 22 22 22 222- R2- 22- 22. 22 :2- 22 22 3:22 :2 22 22 :222-ws2 22 22- .22- .32-222- 22 22 .22 23.8.2.2.” 83 5a 82232 925m ooom 22 $2 22 :2 22 $2 22- .2- 22- 22- 22 22- 22 22- .2... :2- :2 22- a2 22- 22 E2 22- 22- .22- 22- 22. 22- .22- 2:25 :2- 22- 22 22 :2- 22. 22 22 :2 :2- 22 22 2- 2:2 .523 :2 22 22 22- 22 22 22- .22- :22- 22- .2222 $2 $2 23.8.2... 22 22 22 22- 22 22 E2- :2- 22. .22. .22 22. 22 22 2.....E2 .:2 .2252 £22.52 2 2.2- .22- .2222- .2222 .32 .222523. 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Dependent Variable I Independent Vanables ] R2 2000 average stem countsfm ' ii i- ° S. mnericcmus -l 18(Cl) -46(ortho-P) -—264(NO3-N) —3 1(water 0.940 table) +78(conductivity) —105 (alkalinity) M. arvensis 9.9(Cl) —5.8Lalkalinity) 0.349 A. lanceolatus 33.0(alkalinity) 0.229 Mira spp. - -- Other species -11 5(oonduct1v1ty)-15 2(alkal1mty) 0.452 2000 relauve abundance per plot fl S. mericanus -0. 2(0) —0.1(ortho-P) —0.5(NO3-N) 0.908 + .2(conductivity)- .3(alkalinity) M. arvensis 0.02(Alkalinity)+0.03(Cl) 0.389 A. Ianceolatus 0.1(water table) +0.1(ortho—P) 0.388 T who spp. -0. 1(water table) 0.369 Other species -0. l(conduct1v1ty)—0 2(NO3 -N) 0.585 2001 average stem counts/m ..... '* ‘ " "*7: 'i- if S. americanus 54. 4(water table) + 84. 8(pH)-99. 4(ortho-P) 0.895 +120. 3(conductivity) M arvensis +6.1(alkalinity) 0.248 A. lanceolatus -- - T. latzfolia 1.2(conductivit1) + 1.6(ortho-P) 0.494 71 angustg'folia -5.4(oonductivity) 0.292 T min spp. - -- Other species -28. 4(water table)-l 3. 6(alkalm1ty) 0.677 2”! relanve abundance per plot , . " H f.'"575;"7"T‘frgj'f?’5?if?"$5739.32,g5 S. mericanus 0. 1(water table) —0. 2(ortho-P)+0 2(conduct1v1ty) 0.854 M arvensis - A. lanceolatus 0.1(ortho-P) 0.268 71 latrfolia 0.004(water table) +0.009(ortho-P) 0.733 71 angustfi'olia -0.02(conductivity) 0.311 T ”In spp. 0.02(ortho-P)+0.01 (water table) -0.02(conductivity) 0.497 Other species -0.2(water table) -0.06(alkalinity) 0.827 38 S. americanus patch Forested wetland Agricultural Figure la. Location of the Maple River salt seep. The insert shows a map of Michigan with the ¥ indicating the location of the Maple River salt seep in Clinton County, Michigan. 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W m: . . . o m 54 6309.285 m 98 < $85.: 98:3 «8“ >258 demo of mo 5:82 05 83065 :5 “a 353% 2E. .58 mo macaw 05 E 8388 $33 80¢ :83 8355 :08 SO in Ea 832388 BC. .3 Emmi :5 8:33 ON_‘ 03 ow OO Ov om o ON- ow. OO- _ _ _ _ _ _ _ AYo'ulfllnII ‘\nhollu+honnt union: fl-mH6-‘n&ullllw mmcmfiza...o.: ..w b . w 59 0% me 3293 ._ oBmE 2.8 5: Queens: .5 32 58800 a ASSBV 38» x32 ® Assesses 5 :35 O 3555 find 0 58.139533 0 them. Em 78 Figure 2. Unknown Cymbella spp. of benthic diatom algae that was common found in the Dow pond. 79 Chapter 4; Growth of Salt Tflrant mt: in ; Bring 59d Stgrm Water Pond in Miebi an Introduction Environmental conditions strongly influence the composition and structure of a plant community. In ponds and wetlands, plants must deal with the stresses of saturated or inundated soils in which oxygen levels in the root zone are low (Sculthorpe 1967). In salty environments such as along ocean coastlines, plants must deal with high osmotic pressures in the external environment caused by high salt concentrations, in addition to dealing with low oxygen levels in the root zone (Waisel 1972, Queen 1974). Plants have developed several mechanisms to deal with high salt including: succulence (Ungar 1991), accumulation of ions or organic compounds (Broome et al. 1995, Ewing et al. 1989), secretion of salts by salt glands or salt hairs (Waisel 1972), and selective or restrictive ion absorption (Queen 1974, Ungar 1991). In the wetter regions of the midwestern United States, saline wetlands and ponds are historically rare, usually only occurring over fossil salt beds or in the vicinity of salt springs (Waisel 1972). Thus, few native halophytes occur in these areas (Catling and McKay 1981). However, human impacts have increased saline areas in the Midwest by urbanization, road salt runoff, agricultural runoff, and municipal and industrial waste disclmrge (Waisel 1972, Ungar 1974, Williams et al. 1997). The brine pond at the Dow Chemical Company in Midland, Michigan, has been strongly impacted by salinity. Since highly salt impacted areas are relatively rare in Michigan, this pond provided an opportunity to investigate types of plants that could tolerate this habitat after being transplanted into it. 80 The objective of this study was to identify salt tolerant, native Michigan vascular plants that could be used to create wetlands along the margins of the Dow brine pond complex. I assumed that diversifying the plant community would enhance wildlife habitat and encourage use of the pond by a greater diversity of organisms. I hypothesized that native Michigan wetland plants with a high salt tolerance could be used to establish and expand wetland habitat in and around the Dow pond. Four species of emergent wetland plants: common md (Phragmites austmlis), narrow leaf cattail (Typha angusafolia), three square bulrush (Schoenopleclus prmgens), and Olney’s three square bulrush (S. amen'cwms) were used in the transplant study. Mngds and Materials Si ' i n Dow pond - The structure, water chemistry and biota of the Dow brine pond complex are discussed in detail in Chapter 3. An open sandy bank of the main pond was selected for the transplanting experiment (Figure 1). This area had been submerged by pond water in 2000 and prior years according to Dow employees, but water levels had not reached this area in 2001 by the time the rhizome/stem sections of the four selected plant species were transplanted into plots for the experiment. Based on conversations with DOW employees, I assumed that this area would be inundated during the growing season. Unfortunately, this proved to be an incorrect assumption. All growth experiments occurred in the main pond of the Dow brine pond complex (Figure 1). The main pond will be referred to as the “Dow pond” in this chapter. MSU pond - A storm retention pond, called the “MSU pond,” was constructed on the campus of Michigan State University (MSU) to treat runoff from a newly constructed 81 parking lot. The parking lot was constructed across Farm Lane road from the MSU Pavilion south of the Turf Grass Research Facility south of Mt. Hope Road. Runoff from the newly constructed parking lot was routed via a grassy swale into the pond, which was constructed east of the parking lot. In addition, an agricultural tile was intercepted and water from it was routed through the swale and pond to insure that the pond had sufficient water in it to support wetland vegetation. Since the edges of this newly constructed pond consisted of bare sediment, I was able to use the MSU pond as a non- saline reference for the transplanting of three of the emergents planted at the Dow pond (Figure 2). The purpose of this reference was to demonstrate whether these species could successfully be transplanted from the source populations into a fieshwater environment following the same time schedule and procedures used for transplanting these species into the Dow pond. I elected not to transplant the fourth species, Phragmites australis, into this pond given its invasive nature and the desire of project planners to have a more diverse population of native plant species around the MSU pond. Prior to the transplanting experiment, the MSU pond had a heavy bloom of green algae and some small patches of sedges and smartweed growing around the pond edges. Twenty feet or more of bare soil surrounded the pond. Later, grass seed and fertilizer were spread on the bare soil; however, very little grass had grown by the end of the first season. Selection of Salt Tolerant Plants - A literature search revealed salt tolerant wetland plants that grow in Michigan (App. A). In addition, surveys of a natural salt seep community growing along the Maple River in Clinton County, Michigan (see Chapter 2), and the existing plant community at the Dow pond were used to select four species of 82 emergent wetland plants to test in the transplant experiment at the Dow pond. The emergent plants were chosen based on four main criteria: 1) high salt tolerance, 2) availability of source populations, 3) native to Michigan, and 4) usefulness of plant structure to provide wildlife food and habitat. Growth Studies - Two growth experiments were conducted in the Dow main pond using plots cormtructed of topsoil and potted emergent plants placed in shallow water (Figure 1). The four plant species chosen for the emergent plant experiments were S. pungens (three-square bulrush), S. mnericmms(01ney's bulrush), T who migus'tr'folia (narrow leaved cattail), and Phragmites mastralis (common reed). All of these except S. mnen’cmms were already growing in or near the water at the Dow pond. Plugs of S. pungens and T. angustrfolia were obtained from wetlands along Saginaw Bay. S. americanus plugs were obtained from a natural salt seep on the Maple River in Clinton County, Michigan. P. australis plugs were transplanted fiom one of several dense stands growing around the perimeter of the Dow pond. Transplanting the plugs of the four emergent species at the Dow pond did not occur until mid-summer (June 26, 2001 for 71 wrgustr'folia, S prmgens, and P. australis, and June 27, 2001 for S. amerr'camrs). Transplanting stems that late in the season (beyond mid-summer) can prevent successful transplanting (Hammer 1992). To determine whether transplanting could be successfully done this late in the season, plugs of the two bulrushes (June 18 for S. americanus, and June 26 for S. pungens) and II wrgustrfolia (June 21, 2001) were transplanted to plots set up at the MSU pond (Figure 2). The two species of bulrush came fiorn the same source populations as the plants at the Dow pond; however, the T. mrgustIfoIia came from a marsh on the MSU campus. 83 The common reed was not transplanted into the MSU pond, since it is a highly invasive species that spreads easily and is difficult to control (Sculthorpe 1985, Hammer 1992). At the Dow pond, each of the emergent species was transplanted into three replicated experimental plots constructed on shallow areas of the main pond that were exposed during low water conditions (Figures 1, 3 and 4). Topsoil was obtained locally and hauled to the site by Dow contractors. Six inches (15 cm) of topsoil were spread out along the shore in two areas that were expected to be inundated by rising water as the pond was filled during the summer. The nine plots were constructed in a 3 x 27 yd area away from the dense stands of common reed that grew along the shore (Figure 2). Each of the nine plots (three plots allocated to 71 ngusu’folia, three allocated to S. mnericwms, and three allocated to S. pungens) measured 3 x 3 yd. The treatments (species transplanted) were randomly assigned to each of the nine plots. The three plots for the P. australr's were constructed in a 3 x 9 yd area with the same plot dimensions used for the other three species (9 ydz, 6 inches topsoil depth) but were placed adjacent to an existing reed stand (Figure 1). Each plot contained 15 plugs for a total of 45 plugs for each. species. These plugs contained one or more rhizomes and stems of the species being transplanted. The individual plots at the MSU pond were similarly designed to those at the Dow pond (Figure 2 and 3). However, each species was not randomly assigned to plots, but grouped in one area with an empty plot between each planted plot (Figure 2). Since the pond was already covered with topsoil, the plots could be more widely spaced. The plugs were planted at the edge of the pond under the water, and were covered by about 5 cm of water for most of the summer. Based on my understanding of water depths in the Dow pond in previous summers, I assumed that the plots would be inundated within a few weeks of transplanting the plant plugs. However, flooding of the plots did not occur during the 2001 growing season. Since the plots at the Dow pond were not in the pond water, the plots were watered starting July 16 with fresh water at the rate of 0.25-0.5 inches of water/day, 5 days/wk so that the transplants could become established before being exposed to brine from the pond. Then starting in mid-August, the plants were watered 5 days/wk with 0.25-0.5 inches of saline water pumped from the Dow Pond until November 1, 2001. The numbers of surviving and new stems, based on being green in color and/or having new shoots or leaves, were recorded for each plug. Maximum stem lengths were measured periodically from time of transplanting until October 10 at the Dow pond. The number of live plugs was based on the presence of live stems. Stem heights were measured only if they were whole, upright stems (not bent or cut). The stem lengths were not measured on the day they were transplanted, but were measured up to one week later. The number of surviving stems and stem lengths were also recorded periodically at the MSU pond until August 8. Counting of stems was stopped at the MSU pond because the stems became so numerous and spread out that they were difficult to count and made the individual plugs indistinguishable. Toward the end of the summer, a random sample of over 50 plant heights for all three species was taken at the MSU pond to compare to the Dow pond populations. Random samples were taken by tossing a stick into the plot and measuring heights of the nearest three stems. Also, measurements of 85 stem height fiom the S. (men'cams population at the salt seep on the Maple River were randomly sampled to compare to the MSU and Dow pond populations. Pot experiment - Since pond water depth never increased enough to inundate the plots during the summer, a second experiment was initiated in late July with three of the emergent plants (excluding P. australis). Six pots per species with one plant plug per pot wereplaced intheedge ofthe mainpond sothatthebaseoftheplant was inundatedbut the stem extended above the water line. An additional six pots were placed in a wetland pond on the MSU campus as a control. The condition of the transplanted stems (live and growing, dead, senescent) or growth of new stems in each pot was recorded periodically until October 10, 2001. Statistical analyses - Statistical analyses on species specific data were performed using Systat 8.0. Paired Student’s t-tests were used to determine differences between number of live stems per plug and stem lengths per plug for all plots combined on each sampling day within the same population (Dow or MSU). Differences between the Dow and MSU pond populations for the average number of live stems per plug and average stem lengths per plug were determined using a two-group independent t-test. Differences among the three plots within the same population on the same sampling date were determined using ANOVA Finally, the difi‘erent stern lengths between S. americanus populations (Dow, MSU, and Maple River salt seep) were detemrined using a two-group independent t-test. Significant differences were determined at p<0.05. Baum: Water Chemistry - The results of the water chemistry are shown in Table 1. The MSU pond had higher dissolved oxygen and % saturation than the Dow pond, although both were supersaturated. Conductivities were the highest at the Dow pond, followed by the Maple River salt seep, from where the S. wnericarms plant plugs were taken. The MSU pond had the lowest conductivity of all three areas. The Dow pond had very low alkalinity indicating low buffering capacity, but the MSU pond and Maple River seep were comparable. The pH and chloride ion concentration was also highest in the Dow pond, whereas the other two sites were lower. Nitrate concentrations were much higher attheMSUpond, thanattheothersites, butammoniumwas higherattheDowpond. Several ion concentrations were not measured at the MSU pond; however, the Dow pond had much higher ion concentrations than the Maple River salt seep (Table 1). Plot experiment - The results of the plot growth experiments are presented in Figures 5- 19 and Tables 2—4. The results for T. angusnfolia are shown in Figure 5-6. At the Dow pond, this species did send up new shoots while the plots were being watered with fieshwater (57% increase), even though 8 of the 45 plugs appeared dead (Table 2). However, the total number of stems declined after the plots were watered with saline water from the main pond and an additional 6 plugs appeared dead (see Table 2, Figure 5). Average maximum stem length decreased while being watered with fresh water but increased after being watered with pond water (Figure 5). T who angustifolia at the MSU pond significantly increased the total number of living stems by 284% over the summer and only one plug appeared dead (Figure 5, Table 2). Stem lengths decreased over the summer (Figure 6). In comparing the two populations on similar sampling dates, the population at MSU had higher average numbers of living stems initially and at the end of the summer. Initial stern lengths were 87 significantly longer at the MSU pond. Stem lengths were significantly shorter at the Dow pond both after fi’esh watering and salt watering (Table 3 and 4). Results for S. americanus are shown in Figures 7-9. Live stems at the Dow pond increased 16% over the entire summer, even though the number of live plugs decreased by 7 (Table 2). The total number of live stems continued to increase although live plugs decreased by 3 even after the first sampling with pond salt water. Only in the final sampling did total number of live stems decrease and two more plugs appeared dead giving a total of 38 live plugs at the end of the summer (Table 2, Figure 7). Stem lengths also increased over most of the summer after an initial decrease. (Figure 8). At the MSU pond, S amerr'camrs live stems and stem lengths increased over the summer (Figures 7 and 8). This species had the highest growth rate of all the species with an increase of 1444% in total number of live stems over the summer and only one plug appeared dead (Table 2). Initially, the average number of live stems and stem heights were similar for the populations at both ponds (Table 3). Later in the summer, the MSU pond had significantly more live stems and taller stem lengths than at the Dow pond both before and after watering with salt water (Table 4). Comparing the stem lengths to the source population by the Maple River, the MSU pond had similar stem lengths, but the Dow pond had significantly shorter stem lengths both before and after salt watering. Stern lengths were significantly longer at the Dow pond after watering with salt water than before salt watering. (Figure 9). S pungens responded poorly to transplanting at the Dow pond plots. The total number of live stems decreased after the first two samplings, but remained stable the rest of the summer, until it decreased again in the final sampling (Figure 10, Table 2). The number of live plugs decreased greatly with this species, from an initial transplanting of 45 plugs to 7 plugs by the end of the summer (Table 2). Stem lengths decreased significantly alter the first sampling; but began to lengthen after being watered with pond water (Figure l 1). The number of stems increased by 477% over the summer for S. pungens at the MSU pond (Figure 10, Table 4). Average stem lengths maintained the same length except for a small dip in the third sampling (Figure 11). The number of live plugs decreased by four (to 41 live plugs) over the summer (Table 2). The initial average number of live stems was higher at the MSU pond than at the Dow pond, but stem lengths were similar when comparing both populations (Table 3). The number of live stems and the stem lengths were much greater at the MSU pond than at the Dow pond later in the summer (Table 4). Despite being well established in many stands around the Dow pond complex, P. australis responded poorly to transplanting. The total number of live stems decreased greatly after transplanting and decreased by 85% over the summer (Figure 12). The number of live plugs also had a large decrease of 41, only 4 live plugs remained at the end of the summer (Table 2). The average number of live stems decreased even more after being watered with saline pond water (Table 2, Figure 12). The average stem lengths were not significantly different throughout the summer (Figure 13). The average number of live stems per plug in each of the three plots per plant species were not significantly different for most sampling dates. However differences were seen on a few sampling dates for T. angustrfolia at both the Dow and MSU ponds (Fig. 14 and 15, respectively), for S. amen'cmms at the Dow pond (Fig. 16), and for S 89 pungens at the MSU pond (Fig. 17). The only significant differences in average stem lengths per plug for the three plots per plant species was seen in the T. wrgustrfolia plots at the Dow pond (Fig. 18) and in S wnericanus at the MSU pond (Fig. 19). Pot experiment - Because of problems with the transplanting experiment (i.e., pond water not reaching the plots at the Dow pond), a pot growth experiment was set up. This experiment was conducted very late in the growing season (late July). After the first weeks at the Dow pond, above ground stems of all three species appeared dead. However, a week later S. amen'carms had a new shoot appear. After four weeks, S. americanus had two new shoots per pct (12 shoots total) and TI angustzfolia had one new shoot for all six pots. S pungens had no shoots and appeared to be dead. Six pots of all three species were also set up at the MSU pond to see if the transplants would grow so late in the summer. However, animals ate the rhizomes on all the S. pungens and on three of the T. angustrfolia. The pot experiment was run with the remaining three I angusnfolia and six S. americanus pots. Nearly all the original stems appeared dead, but T. angustr‘folia had one new shoot per pot (three total) and S. americanus had an average of four new shoots per pct (24 total). D' . Plot experiments - The plants at the Dow pond grew worse overall than the plants transplanted to the MSU pond in every experiment. The MSU pond is much fiesher than the main pond at Dow (see Table 1). T. angustifolia and S. americams were transplanted 5 days and 9 days earlier, respectively, at the MSU pond than at the Dow pond. Although this may have had some impact on establishment and growth, other factors likely contributed as well to their lower growth such as water levels, salinity and grazing. At the MSU pond, a few centimeters of water covered the plots for nearly the entire summer, whereas water never reached the Dow pond plots without manually watering by a hose. All the species at the Dow pond and T. wrgustzfolia and S. amen'canus at the MSU pond showed none or very few reproductive seed heads. However, S. pungens produced many seed heads in the MSU pond. T angustr‘folia and P. ausr‘ralis were already starting to flower at the time of transplanting. Due to transplanting shoclg above ground shoots often die and new grth fiom shoots occurs, (Hammer 1992) thus the transplants may have put more effort into establishing themselves and vegetatively growing instead of sexually reproducing. The plants at the MSU pond were successful at establishing and greatly proliferated. Although they continued to grow after the early August sampling, they became so numerous it was difficult to count. At the Dow pond, however, few new shoots were added after watering with salt water, but stern length did increase. S. americanus continued growing the first sampling after watering with pond water because the number of live stems increased, but the number of stems decreased in the final sampling at the Dow pond. For both S. prmgens and T. mtgustr‘folia, the number of live stems decreased after watering with salt water. The average maximum stem lengths were significantly taller at the MSU pond than at the Dow pond. In poor environmental conditions, such as at the Dow pond, plants do not grow as well as was seen in this experiment (Begon et al. 1990). At the Dow pond, stem lengths continued to increase after being watered with salt water, although in most cases no new stems were added. 91 The most salt tolerant species was P. australis (Hellings and Gallagher 1992), which was found growing in several areas around the Dow pond perimeter. This species had the worst transplanting record of all four species. As with all the species, they were transplanted late in the season. In temperate regions, the best time to plant is early spring, although planting can occur up until mid-summer (Hammer 1992). Some species, such as cattails and bulrush can be planted later than mid-summer (Hammer 1992), but P. ausnulis does not appear to be as tolerant. S. pungens has the second highest salinity tolerance (App. I), however this species had the second worst growth record at the Dow pond. Although this species is reported to tolerate high salinity, different populations have varying tolerance levels (Ungar 1991). Therefore, the source population of S. pungens may have not had a high salt tolerance. This species' low salt tolerance is also seen in the pot experiment, in which all pots of S prmgens had no new growth, while the other two species produced new shoots. In addition, this species suffered the greatest amount of grazing at the Dow pond. On several occasions, the entire stem and rhizome appeared to be removed. Of all the species, S. amerr'carms grew best at both ponds. S mnericanus had the second lowest salinity tolerance of all the plants tested (App I). This species may have been more successful partly because the phigs had more soil than the other species. The source populations of T. angusnfolia, S pungens and P. australis were all in sandy, loose soils that fell away from the rhizomes upon extraction of the stems. S. amen'canus populations however, were in more clay-like organic soil that held together upon extraction of the plant plugs. Thus, a clump of soil, stems, rhizomes, and possibly seeds were transplanted, whereas the other species only had the stems and rhizomes being 92 transplanted. In addition, the plugs may have prevented the mghly damaging grazing that was done to S pungens by making the rhizomes more difficult to remove. Although the stems were often cut, indicating grazing, the plugs and rhizomes were rarely removed. Even though species with soil plugs grew better, all three species transplanted were successfully established at the MSU pond with only the stem and rhizome. S mnericanus was also successfirl in the pot experiment. This species produced an averageoftwonewshootsperpotattheDowpondwhenitwassaturatedwith salt water. Although it only grew half as well as pots at the MSU pond (producing an average of 4 new stems per pot), it still was able to grow successfully. Thus, this species would be a good candidate to plant at the Dow pond. The least salt tolerant species was T. angustifolia, however, it was the second best grower in the transplant experiment. The highest salt tolerance reported for this species is near 20 ppt (McMillan 1959), which is near the highest salinity reported for the 2001 summer at the Dow pond. T. angustrfolia was abundant on the edges of an adjacent pond, thus as the Dow pond becomes fresher, this species will likely become more abundant at the pond since it is an aggressive invader of open wetlands (Hammer 1992, Prach and Wade 1992). Differences between plots within populations on the same sampling dates were probably mostly affected by wind and grazing. For T. angusrrfolia at the Dow pond, the middle plot consistently had more live stems and taller stem lengths. The middle plot may have been partly protected by the wind by the first plot of T. wigustrfolia that was directly west of the middle plot (Figure 4). The first S mnericams plot at the Dow pond may have been affected by grazing. This plot was between two S pungens plots, which 93 were the most heavily gazed species in this experiment (Figure 4). In addition, this plot had one plug that was completely pulled out by gazers, and it was also the only plot for several samplings that had any dead plugs (2 out of 15). At the MSU pond, the plot farthest east consistently had fewer stems or shorter stems lengths for all three species. This would also indicate that wind my have caused the differences between the plots within this population. Based on the blueprints of the MSUpond, theeast endofthepond is most exposedwithaless steep slopeofahill and no trees. Unfortunately, this transplant study was not continued into a second year since funding fiom Dow was not renewed. Had the study continued, I would have done additional studies to determine if the plants that were transplanted in the first year were well enough established to survive the winter. Tracking the stem counts and stem heights would have been continued into the second year to determine over-winter survival rates. Furthermore, growth studies of these four species should be conducted in the N-S and E-W ponds, which have fresher water. Part of the project goals for the Dow pond was to decrease the salinity of the pond water. Conducting these growth studies in the N- S and E-W ponds could indicate the desired salinity level for the main pond at which successful of transplanting and revegetating of the main pond could occur. Other variations of this transplant study would be to find additional plant species (see App. A) that may tolerate this salinity and conduct similar gowth experiments. An additional study would be to push topsoil into the pond, especially near areas where P. austmlis patches were well established. This plant was growing along the edges, mainly where soil had been pushed into the pond, but it did not appear to be 94 growing on the pond sediments. Although in some areas, P. ausn-aIis had sent horizontal shoots into the pond water, suggesting this species is tolerant of the saline pond water. If P. australis expanded to the topsoil, it would help determine if P. australr's is prohibited fiom gowing in the pond because of the sediments or due to the some other factor. Additionally, the chemistry and engineering groups working on this project suggested that a topsoil cap be placed over the gel-like pond sediments to decrease the diffusion of salt into the pond water in order to freshen the We: This experiment of capping the sediments would be beneficial to both reduce the salt content of the pond water and to help establish plant growth. 95 Works cited Begon, M., J. L. Harper, C. R. Townsend. 1990. Ecology: Individuals, Populations and Communities. 2‘“1 ed. Blackwell Scientific Publications, Boston. Broome, S. W., 1. A. Mendelssohn and K. L. McKee. 1995. Relative growth of Spartina parens (Ait.) Muhl. and Scimus olneyi gray occurring in a mixed stand as affected by salinity and flooding depth. Wetlands 15: 20-30. Catling, P. M. and S. M. McKay. 1981. A review of the occurrence of halophytes in the eastern Great Lakes Region. Michigan Botanist. 20: 167-179. Ewing, K. J. C. Earle, B. Piccinin, and K. A. Kerwhaw. 1989. Vegetation patterns in James Bay coastal marshes. II. Physiological adaptation to salt-induced water stress in three halophytic gaminoids. Can. J. Bot. 67: 521-528. Hammer, D. A 1992. Creating Freshwater Wetlands. Lewis Publishers, Boca Raton. Hellings, S. E. and J. L. Gallagher. 1992. The effects of salinity and flooding on Phragmites australis. Journal of Applied Ecology. 29: 41 -49. McMillan, C. 1959. Salt tolerance within a Twha population. Am. J. Bot. 46: 521-526. Prach, K. and P. M. Wade. 1992. Populations characteristics of expansive perennial herbs. Preslia, Praha. 64: 45-51. Queen, W. H. 1974. Physiology of costal halophytes. In Ecology of Halophytes. Ed R. J. Reimbold and W. H. Queen. Academic Press, New York. Sculthrope, C. D. 1967. The Biology of Aquatic Vascular Plants. Edward Arnold, London. Ungar, I. W. 1974. Inland halophytes of the United States. In Ecology of Halophytes. Ed R. J. Reimbold and W. H. Queen. Academic Press, New York. Ungar, I. W. 1991. Ecophysiology of Vascular Halophytes. CRC Press, Boca Raton. Waisel, Y. 1972. Biology of Halophytes. Academic Press, New York Williams, D. D., N. E. Williams, and Y. Cao. 1997. Spatial differences in macroinvertebrate community structure springs in southeastern Ontario in relation to their chemical and physical environments. Can. J. Zool. 75: 1404-1414. Table 1. Water chemistry data from the Dow pond, MSU pond, and the Maple River salt seep, the source of the S wnericanus population. “n.d.” indicates non-detectible, an “— “ indicates parameters not sampled. Parameter Dow pond MSU pond Maple River Dissolved oxygen (mg/L) 7.8-8.7 10.7-14.8 - % oxygen saturation 92.9-103% 120%-165% - Conductivity (uS) 24,000-38,500 720-747 531-3471 Alkalinity (mg CaC03/L) 67-75 180-194 159-361 PH 8 7.76-7.85 6.98-7.89 N03 —N (mg NIL) 0.13-0.24 12 n.d-7.95 NH4 ~—N (mg N/L) 0.69-0.77 0.0001 1-0.00015 n.d.-.27 Potassium (K, mg/L) 414-441 - n.d-l3.5 Calcium (Ca, mg/L) 5400-5795 - 67-1110 Ortho-Phosphorus (mg/L) n.d. 3.9x10"-4.2x10“ 0.01-0.08 Manganese (Mn, mg/L) 0.2 - 0.02-0.9 Sodium (Na, mg/L) 1728-1874 - 11-666 Chloride (Cl, mg/L) 12,315-14,065 29 68-1800 Table 2. Total number of live stems for all 45 plugs at the initial and final sampling for both the Dow and MSU ponds. Also shown is the total number of live stems before watering with salt water at the Dow pond. The number in parenthesis indicates the number of live plugs, indicated by the presence of living stems. Dow MSU Species Initial Before salt Final Initial Final P. australis 124 (45) 29 (7) l8 (4) - - T. angustr'folia 58 (45) 91 (37) 57 (31) 79 (45) 303 (44) S americanus 307 (45) 413 Q13) 356 (38) 314 (45) 4849 (44) Springer”: 119 (45) 47 (18) 28 (7) 162 (45) 934 (41) Table 3. Initial average stern number per plug and stem lengths per plug with standard deviation at the Dow and MSU ponds with the t and p values. A " indicates a significant difference between populations .05). Species Dow MSU t p 71 angustrfolia # stems 1.29 :tO.458 1.76 10933 2.96 0005* Stem lengths 136 i203 173 i303 6.48 >0.001* S. americcmus # stems 6.82 13.68 6.98 i272 0.25 0.807 Stem lengths 45.8 i242 55.8 $16.14 2.08 0046* S pungens # stems 2.64 i103 3.60 i159 3.54 0001* Stem lengths 60.3 i187 61.9 21:18.9 0.71 0.483 97 Table 4. The average number of stems per plug and stems lengths per plug with standard deviations comparing the Dow and MSU plant populations before (Aug 14) and after (Oct 10) watering with salty pond water at the Dow pond plots with the t and p statistics. All number of stems and stem lengths were significantly different in the two populations (p=0.05). Species Dow MSU t, p Dates Aug 14 Oct 10 Aug 8 Aug 14 Oct 10 T. # stems 2.02 i1.45 1.27 11.07 6.73 2122.93 10.4, 11.5, angustifolia >0.001 >0.001 Stem 42.2 i372 63.7 i182 173 i303 15.6, 9.49, lengths >0.001 >0.001 S # stems 9.17 11:5.83 7.91 i626 108 i387 17.5, 17.4, americanus >0.001 >0.001 Stem 17.4 21:10.6 29.7 :1:15.8 87.1 19.54 31.1, 20.9, lengths >0.001 >0.001 S pungens # stems 1.04 21:1.86 0.622 20.8 i168 7.88, 7.82. i165 >0.001 >0.001 Stem 11.3 i808 34.0 $18.3 64.8 i237 3.31, 0.94, lengths 0.016 0.014 98 Tittabaw ssee River Pump Stati 1 I Pond Main Pond ' l‘ Test plot No. 1 ' 3 plots, 3x9 yd\s‘. . Test Plot No. 3 Test Plot No. 2 I I 3x6 pcts\ 9 plots, 3x27 yds. . OvEr'iBok Pak " ' Figure 1. Experimental plot locations at the brine pond complex showing the four interconnected ponds with the inlet source (*), outlet pump (x), and direction of water flow indicated by the thick black arrows (9) From the pump station pond outlet (x), the water is pumped into the Tittabawassee River. The Raw Brine Pond is not connected to the brine pond complex and was not examined in this study. The dark gray boxes show the locations of the experimental growth plots (test plots no. 1 and 2) and pot experiment (test plot no. 3). ‘. -— ——o - ,__—____,_, - 99 ‘N . T. angustifolia «5‘ 31 11 Figure 2. The MSU pond with location of plots and species planted. Grey plots indicate plots that were planted and white plots indicate empty or unplanted plots. 1 1 t a .1 I r 1 1 100 i C ‘ C (DQKOO 4 Figure 3. The individual plot design for the Dow and MSU ponds showing the locations of each of the 15 plant plugs (black dots) and the spacing (in yards) between each plug. Plot 1 mill:- Plot 2 Twin Plot 3 S. pun Plot 4 S. am Plot 5 S. pun Plot 6 Typha Plot 7 S. am Plot 8 S. pun Plot 9 S. am Figure 4. Plot layout showing the random placement of the T W angusnfolia (Twin), Schoenoplectus pungens (S. pun), and S americanus (S am) at the Dow pond. Plot 1 is facing west and plot 9 is facing east. 10] ' g 290 j 7 1 / :6 190 2 . +MSU . 3 / Salt watenng .3 140 - Fresh water / ' ' D ' 'DO‘” E . 3 3 watering .3 90 j / ,. . ........ a O D. D . . . . ' . . . ..... D '- 40 1 v v v ' Dr D? r T I r I v T I T T ' fig 1 13-Jun 13-Ju1 12-Aug 11-Sep 11-OCI Sampling dates Figure 5. Total number of living stems per sampling date for Twin angustifolia at the MSU and Dow ponds. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. m 2 200 Freshwater £180 watering g 160 3140 -= C as . 3...... g 80 3° \ Sat -{J- Dow 13' 60 8» 40 E 20 2 o 13-Jtn 13-Ju 12-Aug 11Sep 11-Oct Sanpllngdae Figure 6. Average stem lengths per plug for the whole population of T. wrgustifolia at the MSU and Dow ponds. Different capital letters of indicate significant differences in stem lengths between sampling dates at the MSU pond and different lower case letters indicate significant differences in stem length between sampling dates at the Dow pond. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond 102 10000 1 L411) i n 0 z '6 +MSU .. 1000 3 j - - a - -DOW S 5 DD ...... n ......... D. ' - fl 8 ‘1 Do a . "' 4 , Fresh water Salt watering watering 100 ..................... 8-Jun 8-Jul 7-Au9 6-Sep 6-Oct Sampling dates Figure 7. Total number of live stems per sampling date for Schoenoplectus americanus at the MSU and Dow ponds. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. Note: the y-axis is in log scale so differences in the number of stems in the Dow population can be seen more clearly. 1 00 90 80 70 60 50 40 3O 20 1 0 0 18-Jun 18-Jul 17-Aug 16-Sep 16-Oct Sampling date +MSU — 0- Dow Average stem height(cm)lplug Figure 8. Average stem lengths per plug for the whole population of S. americanus at the MSU and Dow ponds. Different capital letters indicate significant differences in stem lengths between sampling dates at the MSU pond and different lower case letters indicate significant differences in stem length between sampling dates at the Dow pond. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. 103 200 r I r r ’E‘ A 8 1. in 150— “ 5 I on C 2 E 1OOL— ’ — 8 m " 7 C m a: B _ a; 50r- 1 .. < 12% O l Saltsee MSU Dow salt p Dow fresh Popumfion Figure 9: Box plot comparing the average stem lengths of S. amerr'canus populations fi'om the Maple River salt seep (Salt seep), the MSU pond (MSU), the Dow pond before being watered with salt water (Dow fresh) and after salt water (Dow salt). The different letters indicate a significant difference between the population’s heights. * indicate outliers in the stem heights. é . a 5 9 1 E 1 '6 f +MSU '- a 3 ; , - - a - - Dow E j . g D‘D ....... a -Q ......... D” ~ . . 1 . ‘ a i :3 Fresh water 33" watenng watering ! 10 1 TI 1 r 1 1 r r r 1 r 1 r r Ifi r r Tfi 18-Jun 1 -Jul 17-Aug 16-Sep 16~Oct Sampling dates Figure 10. Total number of live stems per sampling date for Schoenoplectus pungens at the MSU and Dow ponds. Arrows indicate where watering occurred with fi'esh water and salt water at the Dow pond. Note: the y-axis is in log scale so differences in the number of stems in the Dow population can be seen more clearly. 104 m C a g 70 ABC C E 60 g B A if 50 \ 55° 40 \\ Fresh water 3 +543” '2 E watering , If ‘ - .fi - a - Dow 8 30 b3! 1" aBEcT ' 20 - ’ o g 10 a~‘~~fl’j(3alt watering g be < J a a a a a L a a A A A x a A a a a a a A L aaaaa O 28-Jun 18-Ju1 7-Aug 27-Aug 16-Sep 6-Oct Sampling date Figure 11. Average stem lengths per plug for the whole population of S. pungens at the MSU and Dow ponds. Different capital letters indicate significant differences in stem lengths between sampling dates at the MSU pond and different lower case letters indicate significant differences in stem length between sampling dates at the Dow pond. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. d .5 O A N 0 «Cl § 14.11 1111 on O Fresh water waltering S?“ watering “in ------------- n P O) O J_LIL 1111 .5 0 IL} 0. . ....... .-A Total number of stems N 0 "~43 -------- U 0 o O rrrI—aIrrrry‘filrrrrrTfirrrfrrTrr 18-Jun 8-Jul 28—Jul 17-Aug 6-Sep 26-Sep 16-Oct Sampling date Figure 12. Total number of live stems per sampling date for Phragmites australis at the Dow pond. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. 105 O) O A 01 o o ’l’ 1+4 i I ’ (a) o r 4 1 1 N o H 1 _a O Salt watering 0 r 4 L l } r 1 I r 1 I r r r i r r l r [L r r r r 117 r 28-Jun 18-Jul 07-Aug 27-Aug 16-Sep 06-Oct Sampling dates Average stem length(cm)lplug Figure 13. Average stern lengths per plug for the whole population of P. australis at the Dow pond. No differences were seen in the average lengths per sampling date. Arrows indicate where watering occurred with fresh water and salt water at the Dow pond. 9 8 3 7 I21-Jun g 6 1:122-Jun a. 5 DZ-Jul g 4 I9-Ju1 3 3 1323-Ju1 =2 2 Qéfl 1 o 1 2 3 Plot Figure 14. Average number of live plant stems per plug per plot for T. angustifolia at the MSU pond. Significant differences in number of stems per sampling are indicated by different letters above the bars. 106 3.5 — b a i g 2-5 l26-Jun 3 2 03-Ju1 3. C1 12-Jul 5 1-5 I18-Jul :- 1 — 1:1 14-Aug 0 5 _ Kl 22-Sep ' 12110-Oct o _ 1 2 3 Plot Figure 15. Average number of live plant stems per plug per plot for T. angustifolia for the Dow pond. Significant differences in number of stems per sampling are indicated by different letters above the bars. Solid bars indicate watering with fresh water and striped bars indicate watering with saline pond water. g’ l 3-Ju1 % 12-Jul S g a 18—Jul fi 14-Aug 5’ c1 22-Sep O 3: a 10-Oct 1 2 3 Dow plots Figure 16. Average stem lengths per plug per plot of T. angustifolia for the Dow pond. Significant differences in number of stems per plot per sampling are indicated by different letters above the bars. Solid bars indicate watering with fresh water and striped bars indicate watering with saline pond water. 107 120 - U 3 E 3 l 22-Jun 2’ 2 [:1 2-Ju1 E [:1 9-Jul a I 24-Jul 2 D 11-Aug O > < MSU plots Figure 17. Average stem lengths per plug per plot for S americanus at the MSU pond. Significant differences in number of stems per plot per sampling are indicated by different letters above the bars. 12 - -— T J I 27-Jun 7'9: 10 %T b 1 T [:1 03—Jul a; \ % m 12-Ju1 9- \ ‘ \/ l 18-Jul g $ ’_ E? 1:114-Aug ‘2' § . §¢ [S 22-Sep N '2 §¢ [210-001 1 I 2 3 Plot Figure 18. Average number of live plant stems per plug per plot for S. americanus for the Dow pond. Significant differences in number of stems per sampling are indicated by different letters above the bars. Solid bars indicate watering with fresh water and striped bars indicate watering with saline pond water. 108 # plants per plug l 26-Jun E: 2-Ju1 13 9-Jul l 24-Jul 1:1 8-Aug 2 3 Plot Figure 19. Average stem lengths per plug per plot for S. pungens at the MSU pond. Significant differences in number of stems per plot per sampling are indicated by different letters above the bars. 109 Chapter 5: Conclgsions Salt has played a major role in human physiology and history. It has been used for preserving food, has influenced economics and history, and has affected the environment through natural and anthropogenic processes (Chapter 1). In the environment, many naturally saline environments have been impacted or destroyed by human influences, whereas new saline habitats have been created by human activities. These new saline areas are often a problem because they replace natural fieshwater habitats (i.e. from road salting or sewage), prohibit the intended use of the land (i.e. agiculture and irrigation in arid regions), or are waste products that pose problems for disposal (i.e. from chemical processes in industry). In this thesis, I examined the effects of salts on the plant assemblage of a rare natural salt seep wetland in Michigan (Chapter 2). These seeps result from salt deposit from Paleozoic seas that covered Michigan, but few salt seeps remain. Due to their unique environment, they contain rare and endangered halophytic species, Olney's three- square bulrush (Schoenoplectus mnericanus) and the spike rush, (Eleochrm's parvula), which should be preserved and protected. The abundance of these rare species is correlated with high conductivity and low levels of soluble reactive phosphorus. The second area I examined was the effects of salt on the biota of an unusually saline pond (for the state of Michigan) that was created to contain spent brine at the Dow plant in Midland, Michigan (Chapter 3). The Dow pond had four ponds that were sequentially connected along a salinity gradient from the input area in the N-W pond to intermediate levels of salinity in the E-W pond to the highest levels of salinity in the main and outlet ponds with salinity varying from less than one fourth the salinity of sea water 110 in the N-W pond to two-thirds the salinity of seawater in the main/outlet pond. However, the ion concentrations were different from seawater with less magnesium than seawater with ionic concentrations in decreasing order dominated by chloride, calcium, sodium, and potassium. The diversity of algae, plants, and invertebrates was low in the pond. However, algae, plant and invertebrate diversity decreased along the salinity gradient with highest diversity in the fieshest pond and lowest diversity in the pond with highest salinity. Much of this increase in diversity appeared to be correlated with invasion of the fresher N-S and E-W ponds by Chara, a macroalga, which provides food and cover for invertebrates, waterfowl and other wildlife. Ifdilution of the ponds continues as a consequence of the fieshening of brine well inputs as these wells are remediated, establishment of a more complex and diverse food web may follow. Finally, I conducted experiments on the effects of saline water from the Dow Main Pond on transplant survivorship and growth of four wetland plants (Chapter 4) (S. amerr’canus, S. pungens, Twin angustrfolia, and Phragmites australis). These four emergent plants were transplanted into growth plots at the Dow pond in mid June. Since transplanting occurred later in the season than was ideal, a transplant experiment (excluding P. australis) was set up at a nearly fresh water storm retention pond on the campus of Michigan State University as a reference in order to document that these species could be successfully transplanted at such a late date in the season. Only S. americcmus (Olney's three-square bulrush) increased in number of live stems (by 16%) and showed potential for establishment and gth when irrigated or transplanted into Dow brine pond water. All other species decreased in number of live stems from the initial planting when irrigated or transplanted into brine pond water. In contrast, 111 survivorship and growth was high in the flesh water storm water retention pond at MSU. No species grew as well at the Dow pond as at the MSU pond. All species grew well and more than tripled their original number of live stems at the MSU pond. Problems at the Dow pond included the fact that the plots were not inundated late in the season as I had expected them to be and problems with geese eating the transplanted plants. The fact that S. wnericanus(01ney's three square bulrush) was able to add some shoots under such harsh conditions, suggests that this species may have some promise for establishment if sediments can brought in to cover the margins of the pond and establishment experiments can be conducted so that the plots are inundated soon after transplanting. Experiments to confirm this and that P. ausrralis can be encouraged to colonize such areas by sending rhizomes from existing stands should be the next line of investigation. P. australis is successfully colonizing areas where soil existed or was pushed into the wetlands suggesting that it is also a viable species for providing cover around the edges of the ponds. 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