Fertilization Favors Grasses Over Forbs

Fertilization Favors Grasses Over Forbs | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Fertilization Favors Grasses Over Forbs Gail Chmura, Yiming Wang This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-8319468/v1 This work is licensed under a CC BY 4.0 License Status: Under Review Version 1 posted 5 You are reading this latest preprint version Abstract There are forb-dominated communities unique to salt marshes of the northernmost Atlantic states of the USA, the New Brunswick coast of the Bay of Fundy and the St. Lawrence Estuary that are not associated with exceptionally high soil salinity or saturation. These forb-dominated (predominantly Plantago maritima and Triglochin maritima ) and Spartina patens -dominated communities were subjected to monthly fertilization with a slow-release nitrogen fertilizer in a Bay of Fundy marsh. After 20 years forb cover significantly declined where they had been dominant and are being replaced by grasses (predominantly S. patens and Spartina alterniflora ). This shift is likely due to the increased availability of nitrogen which conferred a competitive advantage to S. patens and S. alterniflora . These grasses require nitrogen for production of osmolytes, chemicals which reduce osmotic stress due to saline soils. In contrast, Plantago maritima and T. maritima do not depend upon nitrogen for their osmolyte production. In marshes to the south forb communities are only associated with exceptionally high soil salinity or saturation. Results of our experiment suggest that absence of forb communities similar to those of higher latitudes could be due to historically greater availability of nitrogen in marshes to the south. nitrogen osmotic stress Plantago maritima Spartina alterniflora Spartina patens Triglochin maritima Figures Figure 1 Figure 2 Introduction In Northwest Atlantic tidal salt marshes graminoids are recognized as the dominant species (Wigand and Roman 2012 , Baldwin et al. 2012 , Pennings et al. 2012 ). Although forbs are common, they are usually reported to occur in low abundance. Widely recognized exceptions are when they are dominant in communities associated with high soil salinity or highly, permanently saturated soils (e.g., Ewanchuk, & Bertness 2004 ). However, on the coasts of the northernmost Atlantic states of the USA, the New Brunswick coast of the Bay of Fundy (Chmura et al. 1997 ) and the St. Lawrence Estuary (Létourneau 1996 ) communities dominated by the forbs Plantago maritima and Triglochin maritima occur outside of those extreme conditions. Controls on the occurrence of those forb communities have been suggested (Chmura et al. 1997 ), but little studied. The controls on occurrence of graminoids in salt marshes of the Northwest Atlantic have been studied, particularly the role of nutrient availability in competitive success of grasses. On the coast of the Northeast USA Levine et al. ( 1998 ) reported that experimental fertilization shifted dominance from Spartina patens (= Sporobolus pumilus ) to Spartina alterniflora (= Sporobolus alterniflorus ) – the two species that comprise most of the vegetation cover there (Wigand and Roman 2012 ). To date, we have not found published studies of the impact of nutrient enrichment on forb-dominated communities. Here we report on the response of a forb community to chronic (long term) fertilization in a tidal salt marsh at Dipper Harbour on the New Brunswick coast of the Bay of Fundy. The response of Plantago maritima , T. maritima and other forbs is compared to that of a S. patens dominated community in the same marsh. Methods Study Site The experiment was carried out in a tidal salt marsh at Dipper Harbour, situated on the New Brunswick coast of the Bay of Fundy (Fig. 1 ). At Dipper Harbour the amplitude of mean tides is 6 m and 8 m for large tides (Canadian Hydrographic Service 1995 ). Vegetation was described by Chmura et al. ( 1997 ). The distribution of graminoids is similar to that in salt marshes along the coast of the Northeast USA. Spartina alterniflora dominant in the lowest vegetated reaches of the marsh, S. patens dominant in the high marsh below J. gerardii , but Puccinellia maritima and Festuca rubra are common, although found in low abundance. Another notable exception is the presence of communities dominated by forbs, primarily Plantago maritima and Triglochin maritima . Forb-dominated assemblages occur not just in pannes, but in better drained soils of the marsh, usually at elevations between S. patens and S. alterniflora. Experiment and Environmental Measurements Permanent 70 cm x 70 cm plots were established haphazardly, a minimum of 3 m apart, in two plant communities. One community was dominated by Plantago maritima and T. maritima , and the other by S. patens . The ecological importance of species was assessed in late August 2005 and 2024, using point intercept (Roman, 2001) to determine the relative cover of species in experimental plots. (As this is a continuing experiment we avoided harvesting plants for biomass.) Treatments consisted of biweekly fertilization during the growing seasons (early May through September) from 2005 to 2007. In 2008, treatments were reduced to monthly applications. Each application consisted of 30 g of Scott’s Turfbuilder, a slow release fertilizer containing 29% N (5.3% ammoniacal N, 13.1% urea-N, 9.7% other water soluble N, and 0.6% water insoluble N); 2% P 2 O 5 , 12% K 2 O, 11% S, 1% water soluble Fe, and 0.5% Mn. When this formulation was no longer available we applied Scott’s Super Turfbuilder, a slow release fertilizer containing 24% N (6.4% ammoniacal N, 9.9% urea-N, 7.1% other water soluble N, and 0.9% water insoluble N); 3% P 2 O 5 , 7% K 2 O, and 7% S. During growing seasons, soil pore water was collected from ≥ 15 cm depth by inserting a perforated plastic tube in the middle of each plot, as described by Yu & Chmura ( 2010 ). When no free pore water was available, the tube was inserted deeper into the soil, up to 30 cm depth. This step enabled retrieval of pore water from most plots, except for, occasionally, some of the Plantago -dominated ones. Salinity of soil pore water was measured using a hand-held refractometer (± 0.1). Plot elevations were measured in 2024 and 2003 with a Sokkia GRX3 Real-Time Kinematic. GRX3 is an all-in-view constellation RTK GNSS receiver applied with the CGVD2013 geoid model, which provides elevation relative to sea level in real time. The post-survey accuracy observed was 0.47 cm in the vertical. Measurements taken at the four corners and the center of each plot were then averaged to represent plot elevation. Statistical Analyses All statistical analyses were performed with IBM SPSS 29. The effect of fertilization on species composition, and elevation change in plots was examined through a T-test. For species competition, the relative cover of forbs and graminoids was analyzed with a general linear model to test the effects of fertilization, salinity, distance from the creek, elevation, and all interactions. Results Environmental Data Average salinity, elevations and distance from plots to the nearest creek are reported in Table 1 while individual plot measurements are reported in Tables 2 and 3. Average salinity of soil pore water was slightly higher in the forb than the S. patens -dominated plots. However, there was no significant difference between control and fertilized plots within the S. patens - or forb-dominated plots. Although plot elevations increased between 2003 and 2024, there was no significant difference between average elevations of fertilized and control plots in either year. The average distance to the nearest creek of the S. patens- dominated plots was more than twice that of the average distance to forb-dominated plots. This distance did not significantly vary between fertilized and control plots of either community. Vegetation By the end of the experiment a total of 13 species were encountered within the experimental plots (Tables 2 and 3). Point intercept measurements included the graminoids Bolboschoenus maritimus , Puccinellia maritima, S. alterniflora , and S. patens as well as the forbs Atriplex patula, Glaux maritima, Limonium carolinianum , Plantago maritima, Argentina anserina (=Potentilla anserina), Salicornia maritima , Spergularia canadensis, Suaeda maritima , and Triglochin maritima . In the originally forb-dominated plots there were notable declines in Plantago maritima and T. maritima , with increases in S. patens , S. alterniflora and S. maritima . Although the overall average species richness of forbs declined under both fertilization and control treatments in the forb plots, these differences were not significant (Table 4). There was even less variability in species richness in the S. patens plots (Table 4). There was no change in the total cover of graminoid and forb species in the S. patens , but significant changes occurred with fertilization of the originally forb-dominated plots (Fig. 2). The average grass cover in the forb-dominated plots increased under both control and fertilization but the increase was not significant. However, under fertilization the forb-dominated plots had a significant reduction in forb cover – by nearly half. Fig. 2 % cover of forbs and graminoids at beginning of experiment and after 20 years of fertilization of a) forb-dominated plots and b) Spartina patens dominated plots. The only significant difference is the decline in forb cover in the originally forb-dominated plots. Neither elevation nor distance from the creek had a significant effect on the decreases in forbs in fertilized forb plots tested by a general linear model. Average soil porewater salinity, which was slightly higher in fertilized forb plots, was identified as a significant factor influencing the forb/ graminoid cover ratio. No interactive effect was observed from these results. Discussion We assume that increased N supplied by our experimental fertilization disproportionally benefited grasses in dealing with salinity stress, allowing them to out-compete forbs. A major mechanism that halophytes employ to deal with the osmotic stress of saline soils is the production of osmolytes. These are compounds that by accumulating in a plant’s cytoplasm balance the osmotic potential of the Na + and Cl − ions that are taken up from the soil and accumulate in vacuoles of the cells (Slama et al 2015 ). There are three primary types of osmolytes quaternary ammonium compounds such as glycine betaine and proline, tertiary sulfonium compounds such as dimethylsulfoniopropionate, and soluble carbohydrates, i.e., sugars and sugar alcohols such as surcrose, mannitol and sorbitol (Jawahar et al. 2019 ). Nitrogen is an important element in both the quaternary ammonium osmolytes and the enzymes that produce them while soluble carbohydrates contain no nitrogen (Cavalieri & Huang 1983 ; Jawahar et al. 2019 ). Hence, increasing the availability of reactive nitrogen confers a major advantage to those species dependent upon glycine betaine and proline. As the grasses S. patens and S. alterniflora are noted for their dependence on proline and glycine betaine (Cavalieri & Huang 1979 . Hester et al. 2001 ) but also the forb Suaeda maritima for proline (Moghaieb et al. 2004 ), their increase in fertilized forb plots is most likely due to their access to increased reactive nitrogen (Cavalieri & Huang 1983 ). We note that salinity was the only environmental characteristic that helped explain the decrease in the forb/graminoid ratio. In contrast, the most important osmolyte used by Plantago maritima is sorbitol (Ahmed et al. 1979 ) while Triglochin maritima seems to use both soluble carbohydrates and proline as osmolytes (Briens & Larher 1982 ). Thus, increased supply of reactive nitrogen through our fertilization would have conferred no advantage to the forbs, particularly with the increasing salinity observed. The exception is S. maritima which did increase, likely linked to its dependence upon glycine betaine and proline as osmolytes (Moghaieb et al. 2004 ). If increased availability of reactive nitrogen favors grasses, we could postulate that the absence of similar forb communities from marshes at lower latitudes on the Northwest Atlantic coast could be related, in part to the historically higher supply of nitrogen there (Galloway et al. 2002 ). The coastline to the south of New Hampshire is highly developed and urbanized while the Bay of Fundy coast is predominantly rural (e.g., Wang et al. 2025 ). The predominance of urban land use would result in high nitrogen loading to estuarine waters (Wang et al. 2022 ; Galloway et al. 2002 ), perhaps conferring the same advantage to S. alterniflora and S. patens that dominate those marshes. Increased nitrogen loading would have begun with land clearance associated with European settlement (Chmura et al. 2004 ) and continue to increase with population growth. To test this theory by reducing nitrogen availability would present major challenges. However, the pollen and macrofossil record found in salt marsh sediments (Beecher & Chmura 2004 ; Daoust et al. 1996 ; Chmura et al. 2001) dating prior to European colonization could reveal prior importance of forb communities like those on the coast of eastern Canada. Declarations Competing interests : The authors have no financial or proprietary interests in any material discussed in this article. Author Contributions: GLC: conceptualization, methodology, investigation, funding acquisition, writing—original draft, and writing—review and editing. YW: investigation, data analyses, review and editing. Acknowledgements We are grateful for the help of numerous field and lab assistants: R. Plant, W. Ampuero-Reyes, K. Nicol, A. Sabourin, G. MacDonald, P. Noel, D. Charbonneau, and M. Khan. We thank the Campbell family for allowing access to the marsh and the Malloch family for their hospitality during field visits. Data Availability: All data is reported within the manuscript. References Ahmed, I., F. Larher, and R. Stewart. 1979. Sorbitol, a compatible osmotic solute in Plantago maritima. New Phytologist 82:671–678. Baldwin, A. H., P. J. Kangas, P. Megonigal, M. C. Perry, and D. F. Whigham. 2012. Chapter 9 Coastal wetlands of Chesapeake Bay. In Wetland habitats of North America ecology and conservation concerns , ed. D. P. Bater, and A. H. Baldwin. 29–43. University of California Press. https://doi.org/10.1525/9780520951419 Beecher, C. B., and G. L. Chmura. 2004. Pollen-vegetation relationships in Bay of Fundy salt marshes. Canadian Journal of Botany 82(5):663–670. Briens, M., and F. Larher. 1982. Osmoregulation in halophytic higher plants: a comparative study of soluble carbohydrates, polyols, betaines and free proline. Plant Cell and Environment 5:287–292. Canadian Hydrographic Service. 1995. Canadian Tide and Current Tables, Volume 1: Atlantic Coast and Bay of Fundy. Ottawa, Ontario, Canada. Cavalieri, A. J., and A. H. C. Huang. 1983. Proline and glycine betaine accumulation by Spartina alterniflora Loisel in response to NaCl and nitrogen in a controlled environment. Oecologia 57:20–24. Cavalieri, A. J., and A. H. C. Huang. 1979. Evaluation of proline accumulation in the adaptation of diverse species of marsh halophytes to the saline environment. American Journal Of Botany 66:307–312. Chmura, G. L., P. Chase, and J. Bercovitch. 1997. Climatic controls on the middle marsh zone in the Bay of Fundy. Estuaries 20:689–699. Chmura, G. L., A. Santos, V. Pospelova, Z. Spasojevic, R. Lam, and J. S. Latimer. 2004. Response of three paleo-primary production proxies to development of an urban estuary. Science of the Total Environment 320:225–243. Daoust, R. J., T. R. Moore, G. L. Chmura, and J. F. Magenheimer. 1996. Chemical evidence of environmental changes and anthropogenic influences in a Bay of Fundy saltmarsh. J Coastal Research 12(2):520–533. Ewanchuk, P. J., and D. Bertness. 2004. Maintenance of high diversity pans in Northern New England salt marshes. Ecology 85:1568–1574. Galloway, J. N., F. J. Dentener, D. G. Capone, E. W. Boyer, R. W. Howarth, S. P. Seitzinger, G. P. Asner, C. C. Cleveland, P. A. Green, E. A. Holland, D. M. Karl, A. F. Michaels, J. H. Porter, A. R. Townsend, and C. J. Vöosmarty. 2002. Nitrogen cycles: past, present, and future. Biogeochemistry 70:153–226. https://doi.org/10.1007/s10533-004-0370-0 Slama, I., C. Abdelly, A. Bouchereau, T. Flowers, and A. Savoure. 2015. Diversity, distribution and roles of osmoprotective compounds accumulated in halophytes under abiotic stress. Annals of Botany 115:433–447. Hester, M. W., I. A. Mendelssohn, and K. L. McKee. 2001. Species and population variation to salinity stress in Panicum hemitomon, Spartina patens, and Spartina alterniflora: morphological and physiological constraints. Environmental and Experimental Botany 46:277–297. Jawahar, G., G. Rajasheker, P. Maheshwari, D. L. Punita, N. Jalaja, Palavalasa Kumari, H. Kumar, S. A. Afreen, R. Karumanchi, A. R. Rathnagiri, and P. Sreenivasulu, N.,Kishor., P.B.K. 2019. Chapter 29 Osmolyte diversity, distribution, and their biosynthetic pathways. In Plant signaling molecules , ed. M.I.R. Khan, P. S. Reddy, A. Ferrante, and N. A. Khan. 449–458. Woodhead Publishing. https://doi.org/10.1016/B978-0-12-816451-8.00028-9 Létourneau, G. 1996. Répertoire des activités de télédétection au Center Saint-Laurent-Module: Marais, maréages et herbiers le long du Saint-Laurent. Environment Canada, Direction de la conservation, Région du Québec. Centre Saint-Laurent Scale 1:20000. Levine, J. M., J. S. Brewer, and M. D. Bertness. 1998. Nutrients, competition and plant zonation in a New England salt marsh. Journal of Ecology 82:285–292. Mansour, M. M. F. 2000. Nitrogen containing compounds and adaptation of plants to salinity stress. Biologia Plantarum 43:491–500. Moghaieb, R. E. A., H. Saneoka, and K. Fujita. 2004. Effect of salinity on osmotic adjustment, glycinebetaine accumulation and the betaine aldehyde dehydrogenase gene expression in two halophytic plants, Salicornia europaea and Suaeda maritima. Plant Science 166:1345–1349. Pennings, S. C., M. Alber, C. R. Alexander, M. Booth, A. Burd, W-J. Cai, C. Craft, C. B. Depratter, D. Di Iorio, C. S. Hopkinson, S. B. Joye, C. D. Meile, W. S. Moore, B. R. Silliman, V. Thompson, and J. P. Wares. 2012. Chapter 4 South Atlantic tidal wetlands. In Wetland habitats of North America ecology and conservation concerns , ed. D. P. Bater, and A. H. Baldwin. 45–61. University of California Press. https://doi.org/10.1525/9780520951419 Roman, C. T., M. J. James-Pirri, and J. F. Heltshe. 2001. Monitoring salt marsh vegetation: a protocol for the long-term coastal ecosystem monitoring program at Cape Cod National Seashore . United States Geological Survey Patuxent Wildlife Research Center. Wang, H., Q. Wang, X. Zhang, S. Du, L. Bai, and S. Xiong. 2025. Mapping global annual urban land cover fractions (2001–2020) derived with multi-objective deep learning. International Journal of Applied Earth Observation and Geoinformation 136. https://doi.org/10.1016/j.jag.2025.104404 Wang, S., Y. Ma, X. Zhang, and Z. Shen. 2022. Transport and sources of nitrogen in stormwater runoff at the urban catchment scale. Science of The Total Environment, 806, Part 1, 150281. https://doi.org/10.1016/j.scitotenv.2021.150281 Wigand, C., and C. T. Roman. 2012. Chapter 2 North Atlantic Coastal Tidal Wetlands. In Wetland habitats of North America ecology and conservation concerns , ed. D. P. Bater, and A. H. Baldwin. 13–28. University of California Press. https://doi.org/10.1525/9780520951419 Yu, O. T., and G. L. Chmura. 2010. Soil carbon may be maintained under grazing in a St Lawrence Estuary tidal marsh. Environmental Conservation 36:312–320. Tables Table 2 Percent cover of species in forb-dominated plots and elevation above sea level (m) at beginning and end of experiment. Soil porewater salinity (PSU) is averaged over experiment and distance to nearest creek (m) measured in 2024. Treatment Control Fertilized Year 2005 2024 2005 2024 Plot 5 6 8 5 6 8 1 3 4 7 1 3 4 7 Elevation above sea level 3.31 3.30 3.25 3.41 3.38 3.35 3.14 3.31 3.28 3.24 3.32 3.51 3.41 3.53 Salinity 29.2 28.3 27.2 26.3 31.3 29.5 28.8 Distance to creek 4.0 3.1 2.2 1.3 5.0 1.0 6.9 graminoids S. patens 0 0 0 39.2 5.1 32.9 0 0 0.7 0 54.8 8.2 8.3 1.3 S. alterniflora 2.3 3.7 1.8 0 3.4 3.9 0.7 11.2 4.1 13.2 45.2 35.3 8.3 37.2 Puccinellia maritima 0 0 0 0 0 11.8 19.0 25.8 9.5 0.0 0 15.3 9.2 17.9 Bolboschoenus maritimus 0 0 0 0 0 0 0 0 0 0 0 3.5 0 0 forbs Plantago maritima 53.9 53.3 64.3 41.8 28.8 43.4 49.0 39.3 47.3 52.9 0 3.5 25.7 5.1 Triglochin maritima 29.7 32.6 25.0 12.7 55.9 6.6 2.7 8.4 29.1 26.5 0 0 0.9 3.8 G. maritima 10.9 9.6 2.7 0 0 0 0.7 9.0 0.7 0 0 0 0 0 L. carolinianum 1.6 0.7 3.6 5.1 6.8 0 0 0 2.0 1.5 0 0 11.9 0 P. anserina 0.8 0 0 0 0 0 0 0 0 0 0 0 0 0 Salicornia maritima 0.8 0 0 0 0 0.0 2.7 0.6 2.0 0 0 0 0.9 0 S. canadensis 0 0 0 0 0 0.0 1.4 0 0 0 0 3.5 11.9 9.0 Suaeda maritima 0 0 2.7 1.3 0 1.3 23.1 4.5 4.7 3.7 0 28.2 22.9 21.8 A. patula 0 0 0 0 0 0 0.7 1.1 0 2.2 0 2.4 0 3.8 Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 16 Dec, 2025 Reviewers invited by journal 11 Dec, 2025 Editor invited by journal 10 Dec, 2025 Editor assigned by journal 09 Dec, 2025 First submitted to journal 09 Dec, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-8319468","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":559010174,"identity":"be405abc-132b-4a99-a2e4-55296f6a963b","order_by":0,"name":"Gail Chmura","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA6ElEQVRIiWNgGAWjYFACxsYDDAwHwKwHQIKHj52wlgaYFmYDkBY2ZiLsgWlhkwCThLTIux8G2lJzJ9rg9uFnVTdqamWAWhg//MCjxfBMIlDLsWe5G86lmd3OOXYc5DBmyR58WhqAWhgbDuduOMNgdju34RhICxsDDz4t/Q9hWti/FcO0MP7B5xcJuC08Zsy5DTVgLcz4bDGQANqScOxw7swzPMXSOccOALUwNkvL4LOlP/3hgw81h3P7zrBv/JxTU2fPz9588OMbfLYcABIJCP5hBlDk4tEAtAVNug6v6lEwCkbBKBiZAAB6sVCUUkMCRAAAAABJRU5ErkJggg==","orcid":"","institution":"McGill University","correspondingAuthor":true,"prefix":"","firstName":"Gail","middleName":"","lastName":"Chmura","suffix":""},{"id":559010175,"identity":"7bae8570-a5af-48f9-aa5c-9bbad1f13606","order_by":1,"name":"Yiming Wang","email":"","orcid":"","institution":"McGill University","correspondingAuthor":false,"prefix":"","firstName":"Yiming","middleName":"","lastName":"Wang","suffix":""}],"badges":[],"createdAt":"2025-12-09 15:39:40","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-8319468/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-8319468/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":100586700,"identity":"df91e4e9-8361-4c7b-b1e7-ca8569b66b26","added_by":"auto","created_at":"2026-01-19 12:07:34","extension":"xml","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":6595,"visible":true,"origin":"","legend":"","description":"","filename":"escoESCOD2500478.xml","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/6c4709f39693cace3de68021.xml"},{"id":100586717,"identity":"0f7910b6-6d08-47b2-ba24-ac5d63454c76","added_by":"auto","created_at":"2026-01-19 12:07:47","extension":"xml","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":982,"visible":true,"origin":"","legend":"","description":"","filename":"ESCOD25004787896.go.xml","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/5574e6de80db3f2f897106c3.xml"},{"id":100586599,"identity":"60d07697-9aa7-4abb-8e6c-305ca276224b","added_by":"auto","created_at":"2026-01-19 12:06:59","extension":"xml","order_by":4,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":729,"visible":true,"origin":"","legend":"","description":"","filename":"ESCOD2500478Import.xml","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/6c0eca3643ab555a2d389275.xml"},{"id":100586759,"identity":"461376f3-8fae-4c6d-85f0-a18ae1d8adfa","added_by":"auto","created_at":"2026-01-19 12:08:05","extension":"xml","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":85862,"visible":true,"origin":"","legend":"","description":"","filename":"ESCOD25004780enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/19186f5be0038336d7c2da9f.xml"},{"id":100586804,"identity":"d8eabda8-a60b-4f8e-b158-72ab1e6458d0","added_by":"auto","created_at":"2026-01-19 12:08:19","extension":"eps","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":135645,"visible":true,"origin":"","legend":"","description":"","filename":"drawingimage1.eps","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/c372ea8246abac9261f25d1f.eps"},{"id":100586685,"identity":"5daf3292-d8d0-49b2-a04b-ed4e2ea79e1e","added_by":"auto","created_at":"2026-01-19 12:07:28","extension":"eps","order_by":7,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":60259,"visible":true,"origin":"","legend":"","description":"","filename":"drawingimage2.eps","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/30dc3c9ecfd647e29ee36a29.eps"},{"id":100586681,"identity":"9486b126-b29e-4e27-97fa-a62d209ca40a","added_by":"auto","created_at":"2026-01-19 12:07:26","extension":"jpeg","order_by":9,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":125475,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage2.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/86b6ca8c722a83c3794f5a1a.jpeg"},{"id":100586697,"identity":"c07b3f12-7617-4a52-bf17-c1c56b6cf803","added_by":"auto","created_at":"2026-01-19 12:07:31","extension":"jpeg","order_by":10,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1074,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage3.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/acc274220013a8e2409a2dc2.jpeg"},{"id":100586673,"identity":"81b18035-93cb-49c9-b6e6-99aafc4c0a74","added_by":"auto","created_at":"2026-01-19 12:07:16","extension":"jpeg","order_by":11,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":407048,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage4.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/eabcf1b17464522c45ea34c9.jpeg"},{"id":100586719,"identity":"af7a333c-3670-418c-9a4a-c3f575dd1b59","added_by":"auto","created_at":"2026-01-19 12:07:48","extension":"jpeg","order_by":12,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":1056220,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage5.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/76a7ff977b408e1b591e1666.jpeg"},{"id":100586667,"identity":"94ff5db5-ec68-490e-839e-4d20b72b8eb5","added_by":"auto","created_at":"2026-01-19 12:07:10","extension":"jpeg","order_by":13,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":560734,"visible":true,"origin":"","legend":"","description":"","filename":"floatimage6.jpeg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/21fb3a7031f9ab94cab1953a.jpeg"},{"id":100586686,"identity":"dfb5e19d-b343-4a99-b77b-16195d99159f","added_by":"auto","created_at":"2026-01-19 12:07:29","extension":"png","order_by":14,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":219179,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/dafbb7734ee51fa08adcd026.png"},{"id":100586716,"identity":"b79c5b43-ae13-4122-a153-10f49e2cbc7c","added_by":"auto","created_at":"2026-01-19 12:07:47","extension":"png","order_by":15,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":24972,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/01785f91d6bbfdf4d692831e.png"},{"id":100586713,"identity":"d5b31363-2af7-4eb0-85de-302dfc7f2050","added_by":"auto","created_at":"2026-01-19 12:07:44","extension":"png","order_by":16,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":935,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/b89fcbde718962088d4324d3.png"},{"id":100586721,"identity":"5b0d06a0-65e1-48aa-90dd-cd019548fc44","added_by":"auto","created_at":"2026-01-19 12:07:49","extension":"png","order_by":17,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":81311,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/b366001b95bdb4d1f8c87bf4.png"},{"id":100586708,"identity":"2e572b96-b8f2-4a45-a1d0-f3c478cdbee3","added_by":"auto","created_at":"2026-01-19 12:07:40","extension":"png","order_by":18,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":207599,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage5.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/ab2d750d9edb6b2316f03a16.png"},{"id":100586729,"identity":"4cf1fd27-47f9-43a1-ba37-0661522ab158","added_by":"auto","created_at":"2026-01-19 12:08:03","extension":"png","order_by":19,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":114171,"visible":true,"origin":"","legend":"","description":"","filename":"Onlinefloatimage6.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/d56fa36d90933ef11fd14b4b.png"},{"id":100586701,"identity":"d2561127-4e02-4acd-92b0-34adbfa377df","added_by":"auto","created_at":"2026-01-19 12:07:35","extension":"xml","order_by":20,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":84616,"visible":true,"origin":"","legend":"","description":"","filename":"ESCOD25004780structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/c0bb07e30896eefbea51a0b7.xml"},{"id":100586707,"identity":"9ac81d22-d356-4b49-bb62-4db3c449c578","added_by":"auto","created_at":"2026-01-19 12:07:39","extension":"html","order_by":21,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":93350,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/364a81395f08070262fa732d.html"},{"id":100586797,"identity":"a72d05d6-cb17-4022-940e-66c5cb2cf91b","added_by":"auto","created_at":"2026-01-19 12:08:12","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":1542371,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of Dipper Harbour (red circle) in context of broader coastal regions. Image Landsat / Copernicus Data SIO, NOAA, US Navy, NGA, GEBCO, ldeo-Coumbia, NSF, NOAA.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/8bdc33dbefdf65293fab363e.png"},{"id":100586815,"identity":"168a6d63-9193-4103-8eec-aa7301589bf2","added_by":"auto","created_at":"2026-01-19 12:08:30","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":53999,"visible":true,"origin":"","legend":"\u003cp\u003e% cover of forbs and graminoids at beginning of experiment and after 20 years of fertilization of a) forb-dominated plots and b) Spartina patens dominated plots. The only significant difference is the decline in forb cover in the originally forb-dominated plots.\u003c/p\u003e","description":"","filename":"1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/2d6b351feb77a1762da0703f.jpg"},{"id":100587326,"identity":"ef1943a4-7217-4596-94df-19fa0ff77f55","added_by":"auto","created_at":"2026-01-19 12:13:51","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2388849,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-8319468/v1/f9aeec88-943e-4894-ba13-34ff6fc9d04f.pdf"}],"financialInterests":"","formattedTitle":"Fertilization Favors Grasses Over Forbs","fulltext":[{"header":"Introduction","content":"\u003cp\u003eIn Northwest Atlantic tidal salt marshes graminoids are recognized as the dominant species (Wigand and Roman \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, Baldwin et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2012\u003c/span\u003e, Pennings et al. \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Although forbs are common, they are usually reported to occur in low abundance. Widely recognized exceptions are when they are dominant in communities associated with high soil salinity or highly, permanently saturated soils (e.g., Ewanchuk, \u0026amp; Bertness \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2004\u003c/span\u003e). However, on the coasts of the northernmost Atlantic states of the USA, the New Brunswick coast of the Bay of Fundy (Chmura et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1997\u003c/span\u003e) and the St. Lawrence Estuary (L\u0026eacute;tourneau \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e1996\u003c/span\u003e) communities dominated by the forbs \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eTriglochin maritima\u003c/em\u003e occur outside of those extreme conditions. Controls on the occurrence of those forb communities have been suggested (Chmura et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), but little studied.\u003c/p\u003e \u003cp\u003eThe controls on occurrence of graminoids in salt marshes of the Northwest Atlantic have been studied, particularly the role of nutrient availability in competitive success of grasses. On the coast of the Northeast USA Levine et al. (\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1998\u003c/span\u003e) reported that experimental fertilization shifted dominance from \u003cem\u003eSpartina patens\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eSporobolus pumilus\u003c/em\u003e) to \u003cem\u003eSpartina alterniflora\u003c/em\u003e (=\u0026thinsp;\u003cem\u003eSporobolus alterniflorus\u003c/em\u003e) \u0026ndash; the two species that comprise most of the vegetation cover there (Wigand and Roman \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). To date, we have not found published studies of the impact of nutrient enrichment on forb-dominated communities.\u003c/p\u003e \u003cp\u003eHere we report on the response of a forb community to chronic (long term) fertilization in a tidal salt marsh at Dipper Harbour on the New Brunswick coast of the Bay of Fundy. The response of \u003cem\u003ePlantago maritima\u003c/em\u003e, \u003cem\u003eT. maritima\u003c/em\u003e and other forbs is compared to that of a \u003cem\u003eS. patens\u003c/em\u003e dominated community in the same marsh.\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy Site\u003c/h2\u003e \u003cp\u003eThe experiment was carried out in a tidal salt marsh at Dipper Harbour, situated on the New Brunswick coast of the Bay of Fundy (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). At Dipper Harbour the amplitude of mean tides is 6 m and 8 m for large tides (Canadian Hydrographic Service \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). Vegetation was described by Chmura et al. (\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). The distribution of graminoids is similar to that in salt marshes along the coast of the Northeast USA. \u003cem\u003eSpartina alterniflora\u003c/em\u003e dominant in the lowest vegetated reaches of the marsh, \u003cem\u003eS. patens\u003c/em\u003e dominant in the high marsh below \u003cem\u003eJ. gerardii\u003c/em\u003e, but \u003cem\u003ePuccinellia maritima\u003c/em\u003e and \u003cem\u003eFestuca rubra\u003c/em\u003e are common, although found in low abundance. Another notable exception is the presence of communities dominated by forbs, primarily \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eTriglochin maritima\u003c/em\u003e. Forb-dominated assemblages occur not just in pannes, but in better drained soils of the marsh, usually at elevations between \u003cem\u003eS. patens\u003c/em\u003e and \u003cem\u003eS. alterniflora.\u003c/em\u003e\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eExperiment and Environmental Measurements\u003c/h3\u003e\n\u003cp\u003ePermanent 70 cm x 70 cm plots were established haphazardly, a minimum of 3 m apart, in two plant communities. One community was dominated by \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eT. maritima\u003c/em\u003e, and the other by \u003cem\u003eS. patens\u003c/em\u003e. The ecological importance of species was assessed in late August 2005 and 2024, using point intercept (Roman, 2001) to determine the relative cover of species in experimental plots. (As this is a continuing experiment we avoided harvesting plants for biomass.)\u003c/p\u003e \u003cp\u003eTreatments consisted of biweekly fertilization during the growing seasons (early May through September) from 2005 to 2007. In 2008, treatments were reduced to monthly applications. Each application consisted of 30 g of Scott\u0026rsquo;s Turfbuilder, a slow release fertilizer containing 29% N (5.3% ammoniacal N, 13.1% urea-N, 9.7% other water soluble N, and 0.6% water insoluble N); 2% P\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e5\u003c/sub\u003e, 12% K\u003csub\u003e2\u003c/sub\u003eO, 11% S, 1% water soluble Fe, and 0.5% Mn. When this formulation was no longer available we applied Scott\u0026rsquo;s Super Turfbuilder, a slow release fertilizer containing 24% N (6.4% ammoniacal N, 9.9% urea-N, 7.1% other water soluble N, and 0.9% water insoluble N); 3% P\u003csub\u003e2\u003c/sub\u003eO\u003csub\u003e5\u003c/sub\u003e, 7% K\u003csub\u003e2\u003c/sub\u003eO, and 7% S.\u003c/p\u003e \u003cp\u003eDuring growing seasons, soil pore water was collected from \u0026ge;\u0026thinsp;15 cm depth by inserting a perforated plastic tube in the middle of each plot, as described by Yu \u0026amp; Chmura (\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). When no free pore water was available, the tube was inserted deeper into the soil, up to 30 cm depth. This step enabled retrieval of pore water from most plots, except for, occasionally, some of the \u003cem\u003ePlantago\u003c/em\u003e-dominated ones. Salinity of soil pore water was measured using a hand-held refractometer (\u0026plusmn;\u0026thinsp;0.1).\u003c/p\u003e \u003cp\u003ePlot elevations were measured in 2024 and 2003 with a Sokkia GRX3 Real-Time Kinematic. GRX3 is an all-in-view constellation RTK GNSS receiver applied with the CGVD2013 geoid model, which provides elevation relative to sea level in real time. The post-survey accuracy observed was 0.47 cm in the vertical. Measurements taken at the four corners and the center of each plot were then averaged to represent plot elevation.\u003c/p\u003e\n\u003ch3\u003eStatistical Analyses\u003c/h3\u003e\n\u003cp\u003eAll statistical analyses were performed with IBM SPSS 29. The effect of fertilization on species composition, and elevation change in plots was examined through a T-test. For species competition, the relative cover of forbs and graminoids was analyzed with a general linear model to test the effects of fertilization, salinity, distance from the creek, elevation, and all interactions.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eEnvironmental Data\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAverage salinity, elevations and distance from plots to the nearest creek are reported in Table 1 while individual plot measurements are reported in Tables 2 and 3. \u0026nbsp; Average salinity of soil pore water was slightly higher in the forb than the \u003cem\u003eS. patens\u003c/em\u003e-dominated plots. However, there was no significant difference between control and fertilized plots within the \u003cem\u003eS. patens\u003c/em\u003e- or forb-dominated plots. \u0026nbsp;Although plot elevations increased between 2003 and 2024, there was no significant difference between average elevations of fertilized and control plots in either year. The average distance to the nearest creek of the \u003cem\u003eS. patens-\u003c/em\u003edominated plots was more than twice that of the average distance to forb-dominated plots. \u0026nbsp;This distance did not significantly vary between fertilized and control plots of either community.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eVegetation\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBy the end of the experiment a total of 13 species were encountered within the experimental plots (Tables 2 and 3). \u0026nbsp;Point intercept measurements included the graminoids \u003cem\u003eBolboschoenus maritimus\u003c/em\u003e, \u003cem\u003ePuccinellia maritima, S. alterniflora\u003c/em\u003e, and \u003cem\u003eS. patens\u003c/em\u003e as well as the forbs \u003cem\u003eAtriplex patula, Glaux maritima, Limonium \u003cem\u003ecarolinianum\u003c/em\u003e, Plantago maritima, \u003cem\u003eArgentina\u003c/em\u003e\u003c/em\u003e \u003cem\u003eanserina\u003c/em\u003e \u003cem\u003e(=Potentilla anserina), Salicornia \u003cem\u003emaritima\u003c/em\u003e, Spergularia canadensis, Suaeda maritima\u003c/em\u003e, and \u003cem\u003eTriglochin maritima\u003c/em\u003e. \u0026nbsp;In the originally forb-dominated plots there were notable declines in \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eT. maritima\u003c/em\u003e, with increases in \u003cem\u003eS. patens\u003c/em\u003e, \u003cem\u003eS. alterniflora\u003c/em\u003e and \u003cem\u003eS. maritima\u003c/em\u003e. \u0026nbsp; Although the overall average species richness of forbs declined under both fertilization and control treatments in the forb plots, these differences were not significant (Table 4). \u0026nbsp;There was even less variability in species richness in the \u003cem\u003eS. patens\u003c/em\u003e plots (Table 4).\u003c/p\u003e\n\u003cp\u003eThere was no change in the total cover of graminoid and forb species in the \u003cem\u003eS. patens\u003c/em\u003e, but significant changes occurred with fertilization of the originally forb-dominated plots (Fig. 2). \u0026nbsp;The average grass cover in the forb-dominated plots increased under both control and fertilization but the increase was not significant. \u0026nbsp;However, under fertilization the forb-dominated plots had a significant reduction in forb cover \u0026ndash; by nearly half.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFig. 2\u003c/strong\u003e % cover of forbs and graminoids at beginning of experiment and after 20 years of fertilization of a) forb-dominated plots and b) Spartina patens dominated plots. The only significant difference is the decline in forb cover in the originally forb-dominated plots.\u003c/p\u003e\n\u003cp\u003eNeither elevation nor distance from the creek had a significant effect on the decreases in forbs in fertilized forb plots tested by a general linear model. Average soil porewater salinity, which was slightly higher in fertilized forb plots, was identified as a significant factor influencing the forb/ graminoid cover ratio. No interactive effect was observed from these results.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eWe assume that increased N supplied by our experimental fertilization disproportionally benefited grasses in dealing with salinity stress, allowing them to out-compete forbs. A major mechanism that halophytes employ to deal with the osmotic stress of saline soils is the production of osmolytes. These are compounds that by accumulating in a plant\u0026rsquo;s cytoplasm balance the osmotic potential of the Na\u003csup\u003e+\u003c/sup\u003e and Cl\u003csup\u003e\u0026minus;\u003c/sup\u003e ions that are taken up from the soil and accumulate in vacuoles of the cells (Slama et al \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). There are three primary types of osmolytes quaternary ammonium compounds such as glycine betaine and proline, tertiary sulfonium compounds such as dimethylsulfoniopropionate, and soluble carbohydrates, i.e., sugars and sugar alcohols such as surcrose, mannitol and sorbitol (Jawahar et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Nitrogen is an important element in both the quaternary ammonium osmolytes and the enzymes that produce them while soluble carbohydrates contain no nitrogen (Cavalieri \u0026amp; Huang \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1983\u003c/span\u003e; Jawahar et al. \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHence, increasing the availability of reactive nitrogen confers a major advantage to those species dependent upon glycine betaine and proline. As the grasses \u003cem\u003eS. patens\u003c/em\u003e and \u003cem\u003eS. alterniflora\u003c/em\u003e are noted for their dependence on proline and glycine betaine (Cavalieri \u0026amp; Huang \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1979\u003c/span\u003e. Hester et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2001\u003c/span\u003e) but also the forb \u003cem\u003eSuaeda maritima\u003c/em\u003e for proline (Moghaieb et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2004\u003c/span\u003e), their increase in fertilized forb plots is most likely due to their access to increased reactive nitrogen (Cavalieri \u0026amp; Huang \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1983\u003c/span\u003e). We note that salinity was the only environmental characteristic that helped explain the decrease in the forb/graminoid ratio.\u003c/p\u003e \u003cp\u003eIn contrast, the most important osmolyte used by \u003cem\u003ePlantago maritima\u003c/em\u003e is sorbitol (Ahmed et al. \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1979\u003c/span\u003e) while \u003cem\u003eTriglochin maritima\u003c/em\u003e seems to use both soluble carbohydrates and proline as osmolytes (Briens \u0026amp; Larher \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1982\u003c/span\u003e). Thus, increased supply of reactive nitrogen through our fertilization would have conferred no advantage to the forbs, particularly with the increasing salinity observed. The exception is \u003cem\u003eS. maritima\u003c/em\u003e which did increase, likely linked to its dependence upon glycine betaine and proline as osmolytes (Moghaieb et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2004\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eIf increased availability of reactive nitrogen favors grasses, we could postulate that the absence of similar forb communities from marshes at lower latitudes on the Northwest Atlantic coast could be related, in part to the historically higher supply of nitrogen there (Galloway et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). The coastline to the south of New Hampshire is highly developed and urbanized while the Bay of Fundy coast is predominantly rural (e.g., Wang et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). The predominance of urban land use would result in high nitrogen loading to estuarine waters (Wang et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2022\u003c/span\u003e; Galloway et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2002\u003c/span\u003e), perhaps conferring the same advantage to \u003cem\u003eS. alterniflora\u003c/em\u003e and \u003cem\u003eS. patens\u003c/em\u003e that dominate those marshes. Increased nitrogen loading would have begun with land clearance associated with European settlement (Chmura et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2004\u003c/span\u003e) and continue to increase with population growth. To test this theory by reducing nitrogen availability would present major challenges. However, the pollen and macrofossil record found in salt marsh sediments (Beecher \u0026amp; Chmura \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Daoust et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Chmura et al. 2001) dating prior to European colonization could reveal prior importance of forb communities like those on the coast of eastern Canada.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e \u003ch2\u003eCompeting interests\u003c/b\u003e:\u003c/h2\u003e \u003cp\u003eThe authors have no financial or proprietary interests in any material discussed in this article.\u003c/p\u003e \u003c/p\u003e\u003ch2\u003eAuthor Contributions:\u003c/h2\u003e \u003cp\u003eGLC: conceptualization, methodology, investigation, funding acquisition, writing\u0026mdash;original draft, and writing\u0026mdash;review and editing. YW: investigation, data analyses, review and editing.\u003c/p\u003e\u003ch2\u003eAcknowledgements\u003c/h2\u003e \u003cp\u003eWe are grateful for the help of numerous field and lab assistants: R. Plant, W. Ampuero-Reyes, K. Nicol, A. Sabourin, G. MacDonald, P. Noel, D. Charbonneau, and M. Khan. We thank the Campbell family for allowing access to the marsh and the Malloch family for their hospitality during field visits.\u003c/p\u003e\u003ch2\u003eData Availability:\u003c/h2\u003e \u003cp\u003eAll data is reported within the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAhmed, I., F. Larher, and R. Stewart. 1979. Sorbitol, a compatible osmotic solute in Plantago maritima. \u003cem\u003eNew Phytologist\u003c/em\u003e 82:671\u0026ndash;678.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBaldwin, A. H., P. J. Kangas, P. Megonigal, M. C. Perry, and D. F. Whigham. 2012. Chapter 9 Coastal wetlands of Chesapeake Bay. In \u003cem\u003eWetland habitats of North America ecology and conservation concerns\u003c/em\u003e, ed. D. P. Bater, and A. H. Baldwin. 29\u0026ndash;43. University of California Press. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1525/9780520951419\u003c/span\u003e\u003cspan address=\"10.1525/9780520951419\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBeecher, C. B., and G. L. Chmura. 2004. Pollen-vegetation relationships in Bay of Fundy salt marshes. \u003cem\u003eCanadian Journal of Botany\u003c/em\u003e 82(5):663\u0026ndash;670.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBriens, M., and F. Larher. 1982. Osmoregulation in halophytic higher plants: a comparative study of soluble carbohydrates, polyols, betaines and free proline. \u003cem\u003ePlant Cell and Environment\u003c/em\u003e 5:287\u0026ndash;292.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCanadian Hydrographic Service. 1995. Canadian Tide and Current Tables, Volume 1: Atlantic Coast and Bay of Fundy. Ottawa, Ontario, Canada.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCavalieri, A. J., and A. H. C. Huang. 1983. Proline and glycine betaine accumulation by \u003cem\u003eSpartina alterniflora\u003c/em\u003e Loisel in response to NaCl and nitrogen in a controlled environment. \u003cem\u003eOecologia\u003c/em\u003e 57:20\u0026ndash;24.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCavalieri, A. J., and A. H. C. Huang. 1979. Evaluation of proline accumulation in the adaptation of diverse species of marsh halophytes to the saline environment. \u003cem\u003eAmerican Journal Of Botany\u003c/em\u003e 66:307\u0026ndash;312.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChmura, G. L., P. Chase, and J. Bercovitch. 1997. Climatic controls on the middle marsh zone in the Bay of Fundy. \u003cem\u003eEstuaries\u003c/em\u003e 20:689\u0026ndash;699.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChmura, G. L., A. Santos, V. Pospelova, Z. Spasojevic, R. Lam, and J. S. Latimer. 2004. Response of three paleo-primary production proxies to development of an urban estuary. \u003cem\u003eScience of the Total Environment\u003c/em\u003e 320:225\u0026ndash;243.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDaoust, R. J., T. R. Moore, G. L. Chmura, and J. F. Magenheimer. 1996. Chemical evidence of environmental changes and anthropogenic influences in a Bay of Fundy saltmarsh. \u003cem\u003eJ Coastal Research\u003c/em\u003e 12(2):520\u0026ndash;533.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eEwanchuk, P. J., and D. Bertness. 2004. Maintenance of high diversity pans in Northern New England salt marshes. \u003cem\u003eEcology\u003c/em\u003e 85:1568\u0026ndash;1574.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGalloway, J. N., F. J. Dentener, D. G. Capone, E. W. Boyer, R. W. Howarth, S. P. Seitzinger, G. P. Asner, C. C. Cleveland, P. A. Green, E. A. Holland, D. M. Karl, A. F. Michaels, J. H. Porter, A. R. Townsend, and C. J. V\u0026ouml;osmarty. 2002. Nitrogen cycles: past, present, and future. \u003cem\u003eBiogeochemistry\u003c/em\u003e 70:153\u0026ndash;226. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1007/s10533-004-0370-0\u003c/span\u003e\u003cspan address=\"10.1007/s10533-004-0370-0\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSlama, I., C. Abdelly, A. Bouchereau, T. Flowers, and A. Savoure. 2015. Diversity, distribution and roles of osmoprotective compounds accumulated in halophytes under abiotic stress. \u003cem\u003eAnnals of Botany\u003c/em\u003e 115:433\u0026ndash;447.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHester, M. W., I. A. Mendelssohn, and K. L. McKee. 2001. Species and population variation to salinity stress in Panicum hemitomon, Spartina patens, and Spartina alterniflora: morphological and physiological constraints. \u003cem\u003eEnvironmental and Experimental Botany\u003c/em\u003e 46:277\u0026ndash;297.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJawahar, G., G. Rajasheker, P. Maheshwari, D. L. Punita, N. Jalaja, Palavalasa Kumari, H. Kumar, S. A. Afreen, R. Karumanchi, A. R. Rathnagiri, and P. Sreenivasulu, N.,Kishor., P.B.K. 2019. Chapter 29 Osmolyte diversity, distribution, and their biosynthetic pathways. In \u003cem\u003ePlant signaling molecules\u003c/em\u003e, ed. M.I.R. Khan, P. S. Reddy, A. Ferrante, and N. A. Khan. 449\u0026ndash;458. Woodhead Publishing. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/B978-0-12-816451-8.00028-9\u003c/span\u003e\u003cspan address=\"10.1016/B978-0-12-816451-8.00028-9\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eL\u0026eacute;tourneau, G. 1996. R\u0026eacute;pertoire des activit\u0026eacute;s de t\u0026eacute;l\u0026eacute;d\u0026eacute;tection au Center Saint-Laurent-Module: Marais, mar\u0026eacute;ages et herbiers le long du Saint-Laurent. Environment Canada, Direction de la conservation, R\u0026eacute;gion du Qu\u0026eacute;bec. \u003cem\u003eCentre Saint-Laurent Scale\u003c/em\u003e 1:20000.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLevine, J. M., J. S. Brewer, and M. D. Bertness. 1998. Nutrients, competition and plant zonation in a New England salt marsh. \u003cem\u003eJournal of Ecology\u003c/em\u003e 82:285\u0026ndash;292.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMansour, M. M. F. 2000. Nitrogen containing compounds and adaptation of plants to salinity stress. \u003cem\u003eBiologia Plantarum\u003c/em\u003e 43:491\u0026ndash;500.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMoghaieb, R. E. A., H. Saneoka, and K. Fujita. 2004. Effect of salinity on osmotic adjustment, glycinebetaine accumulation and the betaine aldehyde dehydrogenase gene expression in two halophytic plants, Salicornia europaea and Suaeda maritima. \u003cem\u003ePlant Science\u003c/em\u003e 166:1345\u0026ndash;1349.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePennings, S. C., M. Alber, C. R. Alexander, M. Booth, A. Burd, W-J. Cai, C. Craft, C. B. Depratter, D. Di Iorio, C. S. Hopkinson, S. B. Joye, C. D. Meile, W. S. Moore, B. R. Silliman, V. Thompson, and J. P. Wares. 2012. Chapter 4 South Atlantic tidal wetlands. In \u003cem\u003eWetland habitats of North America ecology and conservation concerns\u003c/em\u003e, ed. D. P. Bater, and A. H. Baldwin. 45\u0026ndash;61. University of California Press. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1525/9780520951419\u003c/span\u003e\u003cspan address=\"10.1525/9780520951419\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRoman, C. T., M. J. James-Pirri, and J. F. Heltshe. 2001. \u003cem\u003eMonitoring salt marsh vegetation: a protocol for the long-term coastal ecosystem monitoring program at Cape Cod National Seashore\u003c/em\u003e. United States Geological Survey Patuxent Wildlife Research Center.\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang, H., Q. Wang, X. Zhang, S. Du, L. Bai, and S. Xiong. 2025. Mapping global annual urban land cover fractions (2001\u0026ndash;2020) derived with multi-objective deep learning. \u003cem\u003eInternational Journal of Applied Earth Observation and Geoinformation\u003c/em\u003e 136. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.jag.2025.104404\u003c/span\u003e\u003cspan address=\"10.1016/j.jag.2025.104404\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang, S., Y. Ma, X. Zhang, and Z. Shen. 2022. Transport and sources of nitrogen in stormwater runoff at the urban catchment scale. Science of The Total Environment, 806, Part 1, 150281. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1016/j.scitotenv.2021.150281\u003c/span\u003e\u003cspan address=\"10.1016/j.scitotenv.2021.150281\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWigand, C., and C. T. Roman. 2012. Chapter 2 North Atlantic Coastal Tidal Wetlands. In \u003cem\u003eWetland habitats of North America ecology and conservation concerns\u003c/em\u003e, ed. D. P. Bater, and A. H. Baldwin. 13\u0026ndash;28. University of California Press. \u003cspan class=\"ExternalRef\"\u003e\u003cspan class=\"RefSource\"\u003ehttps://doi.org/10.1525/9780520951419\u003c/span\u003e\u003cspan address=\"10.1525/9780520951419\" targettype=\"DOI\" class=\"RefTarget\"\u003e\u003c/span\u003e\u003c/span\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYu, O. T., and G. L. Chmura. 2010. Soil carbon may be maintained under grazing in a St Lawrence Estuary tidal marsh. \u003cem\u003eEnvironmental Conservation\u003c/em\u003e 36:312\u0026ndash;320.\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/69519_bce2c0439cd956a6/69519_custom_files/img1768823723.png\"\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"836\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"18\" valign=\"bottom\" style=\"width: 836px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e Percent cover of species in forb-dominated plots and elevation above sea level (m) at beginning and end of experiment. Soil porewater salinity (PSU) is averaged over experiment and distance to nearest creek (m) measured in 2024.\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eTreatment\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"7\" valign=\"bottom\" style=\"width: 273px;\"\u003e\n \u003cp\u003eControl\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"9\" valign=\"bottom\" style=\"width: 358px;\"\u003e\n \u003cp\u003eFertilized\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eYear\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"bottom\" style=\"width: 127px;\"\u003e\n \u003cp\u003e2005\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"3\" valign=\"bottom\" style=\"width: 127px;\"\u003e\n \u003cp\u003e2024\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"bottom\" style=\"width: 170px;\"\u003e\n \u003cp\u003e2005\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"4\" valign=\"bottom\" style=\"width: 170px;\"\u003e\n \u003cp\u003e2024\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003ePlot\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e7\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eElevation above sea level\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.30\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.25\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.35\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.14\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.31\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.24\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.32\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.41\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.53\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eSalinity\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e29.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e28.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e27.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e26.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e31.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e29.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e28.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eDistance to creek\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e6.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003egraminoids\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eS. patens\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e39.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e32.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e54.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eS. alterniflora\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e11.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e13.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e45.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e35.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e37.2\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003ePuccinellia maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e11.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e19.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e25.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e9.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e15.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e9.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e17.9\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eBolboschoenus maritimus\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003eforbs\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003ePlantago maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e53.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e53.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e64.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e41.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e28.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e43.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e49.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e39.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e47.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e52.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e25.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eTriglochin maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e29.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e32.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e25.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e12.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e55.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e6.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e8.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e29.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e26.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eG. maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e10.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e9.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e9.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eL. carolinianum\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e5.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e6.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e11.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eP. anserina\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eSalicornia maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.8\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.6\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eS. canadensis\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e11.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e9.0\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eSuaeda maritima\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.3\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e23.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4.5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e4.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e28.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e22.9\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e21.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"bottom\" style=\"width: 186px;\"\u003e\n \u003cp\u003e\u003cem\u003eA. patula\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 20px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0.7\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e1.1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.2\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 18px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e2.4\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 42px;\"\u003e\n \u003cp\u003e3.8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/69519_bce2c0439cd956a6/69519_custom_files/img1768823748.png\"\u003e\u003c/p\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003e\u003cimg src=\"https://myfiles.space/user_files/69519_bce2c0439cd956a6/69519_custom_files/img1768823760.png\"\u003e\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":true,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"estuaries-and-coasts","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"esco","sideBox":"Learn more about [Estuaries and Coasts](https://www.springer.com/journal/12237)","snPcode":"12237","submissionUrl":"https://www.editorialmanager.com/esco/","title":"Estuaries and Coasts","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"nitrogen, osmotic stress, Plantago maritima, Spartina alterniflora, Spartina patens, Triglochin maritima","lastPublishedDoi":"10.21203/rs.3.rs-8319468/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-8319468/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThere are forb-dominated communities unique to salt marshes of the northernmost Atlantic states of the USA, the New Brunswick coast of the Bay of Fundy and the St. Lawrence Estuary that are not associated with exceptionally high soil salinity or saturation. These forb-dominated (predominantly \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eTriglochin maritima\u003c/em\u003e) and \u003cem\u003eSpartina patens\u003c/em\u003e-dominated communities were subjected to monthly fertilization with a slow-release nitrogen fertilizer in a Bay of Fundy marsh. After 20 years forb cover significantly declined where they had been dominant and are being replaced by grasses (predominantly \u003cem\u003eS. patens\u003c/em\u003e and \u003cem\u003eSpartina alterniflora\u003c/em\u003e). This shift is likely due to the increased availability of nitrogen which conferred a competitive advantage to \u003cem\u003eS. patens\u003c/em\u003e and \u003cem\u003eS. alterniflora\u003c/em\u003e. These grasses require nitrogen for production of osmolytes, chemicals which reduce osmotic stress due to saline soils. In contrast, \u003cem\u003ePlantago maritima\u003c/em\u003e and \u003cem\u003eT. maritima\u003c/em\u003e do not depend upon nitrogen for their osmolyte production. In marshes to the south forb communities are only associated with exceptionally high soil salinity or saturation. Results of our experiment suggest that absence of forb communities similar to those of higher latitudes could be due to historically greater availability of nitrogen in marshes to the south.\u003c/p\u003e","manuscriptTitle":"Fertilization Favors Grasses Over Forbs","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-19 12:01:13","doi":"10.21203/rs.3.rs-8319468/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-12-16T12:05:10+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-12-11T18:54:38+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"Estuaries and Coasts","date":"2025-12-10T23:20:56+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-12-09T16:22:37+00:00","index":"","fulltext":""},{"type":"submitted","content":"Estuaries and Coasts","date":"2025-12-09T10:39:09+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"estuaries-and-coasts","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"esco","sideBox":"Learn more about [Estuaries and Coasts](https://www.springer.com/journal/12237)","snPcode":"12237","submissionUrl":"https://www.editorialmanager.com/esco/","title":"Estuaries and Coasts","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"88ea3a07-446c-4055-a9e1-037abb116c41","owner":[],"postedDate":"January 19th, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2026-03-23T15:14:19+00:00","versionOfRecord":[],"versionCreatedAt":"2026-01-19 12:01:13","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-8319468","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-8319468","identity":"rs-8319468","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}