Register      Login
Marine and Freshwater Research Marine and Freshwater Research Society
Advances in the aquatic sciences
RESEARCH ARTICLE

Why has Phragmites australis persisted in the increasingly saline Gippsland Lakes? A test of three competing hypotheses

Paul I. Boon A F , Doug Frood B , Alison Oates C , Jim Reside D and Neville Rosengren E
+ Author Affiliations
- Author Affiliations

A School of Geography, The University of Melbourne, 221 Bouverie Street, Carlton, Vic. 3053, Australia.

B Pathways Bushland & Environment, PO Box 360, Greensborough, Vic. 3088, Australia.

C Oates Environmental Consulting, 2/44 School Avenue, Newhaven, Vic. 3925, Australia.

D Wildlife Unlimited, PO Box 255, Bairnsdale, Vic. 3875, Australia.

E School of Life Sciences, La Trobe University, Bundoora, Vic. 3086, Australia.

F Corresponding author. Email: paul.boon@unimelb.edu.au

Marine and Freshwater Research 70(4) 469-492 https://doi.org/10.1071/MF18145
Submitted: 4 April 2018  Accepted: 23 July 2018   Published: 9 November 2018

Abstract

Common reed Phragmites australis is the dominant vascular plant species of the shorelines of the Gippsland Lakes, south-eastern Australia. Although substantial declines have been reported for over 50 years, with increasing salinity posited as the cause, P. australis still occurs around the Gippsland Lakes, including in environments with near-oceanic salinities. The occurrence of P. australis in highly saline environments cannot be explained in terms of either seasonal variations in surface water salinity or a freshwater subsidy provided by intrusions of non-saline groundwater into the root zone. An experimental growth trial with plants of different provenance showed that P. australis grew vigorously even at 8–16 PSU (with maximum aboveground biomass at 2–4 PSU). There was some evidence that specimens from saltier sites were more salt tolerant than those from fresher sites. The selection of salt-tolerant strains is the most likely explanation for the occurrence of P. australis in saline sites. However, anthropogenic salinisation is unlikely to be the only factor involved in the historical loss of reed beds, and lower and more stable water levels following the permanent opening of the Gippsland Lakes to the ocean in 1889 are probably also contributing factors.


References

Achenbach, L., and Brix, H. (2014). Can differences in salinity tolerance explain the distribution of four genetically distinct lineages of Phragmites australis in the Mississippi River delta? Hydrobiologia 737, 5–23.
Can differences in salinity tolerance explain the distribution of four genetically distinct lineages of Phragmites australis in the Mississippi River delta?Crossref | GoogleScholarGoogle Scholar |

Achenbach, L., Eller, F., Nguyen, L. X., and Brix, H. (2013). Differences in salinity tolerance of genetically distinct Phragmites australis clones. AoB Plants 5, plt019.
Differences in salinity tolerance of genetically distinct Phragmites australis clones.Crossref | GoogleScholarGoogle Scholar |

Adams, J. B., and Bate, G. C. (1999). Growth and photosynthetic performance of Phragmites australis in estuarine waters: a field and experimental evaluation. Aquatic Botany 64, 359–367.
Growth and photosynthetic performance of Phragmites australis in estuarine waters: a field and experimental evaluation.Crossref | GoogleScholarGoogle Scholar |

Adams, J. B., Veldkornet, D., and Tabot, P. (2016). Distribution of macrophyte species and habitats in South African estuaries. South African Journal of Botany 107, 5–11.
Distribution of macrophyte species and habitats in South African estuaries.Crossref | GoogleScholarGoogle Scholar |

Agriculture Victoria (1999). Measuring the salinity of water. Available at http://agriculture.vic.gov.au/agriculture/farm-management/soil-and-water/salinity/measuring-the-salinity-of-water [Verified 1 August 2017].

Alvarez, M. G., Tron, F., and Mauchamp, A. (2005). Sexual versus asexual colonization by Phragmites australis: 25-year reed dynamics in a Mediterranean marsh, southern France. Wetlands 25, 639–647.
Sexual versus asexual colonization by Phragmites australis: 25-year reed dynamics in a Mediterranean marsh, southern France.Crossref | GoogleScholarGoogle Scholar |

Barrett-Lennard, E. G. (2003). The interaction between waterlogging and salinity in higher plants: causes, consequences and implications. Plant and Soil 253, 35–54.
The interaction between waterlogging and salinity in higher plants: causes, consequences and implications.Crossref | GoogleScholarGoogle Scholar |

Bart, D., and Hartman, J. M. (2003). The role of large rhizome dispersal and low salinity windows in the establishment of common reed, Phragmites australis, in salt marshes: new links to human activities. Estuaries 26, 436–443.
The role of large rhizome dispersal and low salinity windows in the establishment of common reed, Phragmites australis, in salt marshes: new links to human activities.Crossref | GoogleScholarGoogle Scholar |

Batriu, E., Ninot, J. M., and Pino, J. (2015). Interactions between transplants of Phragmites australis and Juncus acutus in Mediterranean coastal marshes: the modulating role of environmental gradients. Aquatic Botany 124, 29–38.
Interactions between transplants of Phragmites australis and Juncus acutus in Mediterranean coastal marshes: the modulating role of environmental gradients.Crossref | GoogleScholarGoogle Scholar |

Bird, E. C. F. (1961a). Landform changes at Lakes Entrance. Victorian Naturalist 77, 137–146.

Bird, E. C. F. (1961b). Reed growth in the Gippsland Lakes. Victorian Naturalist 77, 262–268.

Bird, E. C. F. (1962). The river deltas of the Gippsland Lakes. Proceedings of the Royal Society of Victoria 75, 65–74.

Bird, E. C. F. (1965). ‘A Geomorphological Study of the Gippsland Lakes.’ (Australian National University: Canberra, ACT, Australia.)

Bird, E. C. F. (1966). The impact of man on the Gippsland Lakes, Australia. In ‘Geography as Human Ecology. Methodology by Example’. (Eds S. R. Eyre and G. R. J. Jones.) pp. 55–73. (Edward Arnold: London, UK.)

Bird, E. C. F. (1970). The Mitchell River silt jetties. Victorian Naturalist 87, 162–168.

Bird, E. C. F. (1983). Shoreline changes in the Gippsland Lakes 1957–1983. Proceedings of the Royal Society of Victoria 95, 227–235.

Bird, E. C. F. and Lennon, J. (1989). ‘Making an Entrance: The Story of the Artificial Entrance to the Gippsland Lakes.’ (Geostudies Australia: Bairnsdale, Vic., Australia.)

Bird, E. C. F., and Rosengren, N. J. (1971). The disappearing Mitchell delta. Proceedings of the Royal Society of Victoria 84, 153–158.

Blanch, S. J., Ganf, G. G., and Walker, K. F. (1999). Tolerance of riverine plants to flooding and exposure indicated by water regime. Regulated Rivers: Research and Management 15, 43–62.
Tolerance of riverine plants to flooding and exposure indicated by water regime.Crossref | GoogleScholarGoogle Scholar |

Blanch, S. J., Ganf, G. G., and Walker, K. F. (2000). Water regimes and littoral plants in four weir pools of the River Murray, Australia. Regulated Rivers: Research and Management 16, 445–456.
Water regimes and littoral plants in four weir pools of the River Murray, Australia.Crossref | GoogleScholarGoogle Scholar |

Boar, R. R., Crook, C. E., and Moss, B. (1989). Regression of Phragmites australis reedswamps and recent changes of water chemistry in the Norfolk Broadlands, England. Aquatic Botany 35, 41–55.
Regression of Phragmites australis reedswamps and recent changes of water chemistry in the Norfolk Broadlands, England.Crossref | GoogleScholarGoogle Scholar |

Boon, P. I., Raulings, E., Roache, M., and Morris, K. (2008). Vegetation changes over a four-decade period in Dowd Morass, a brackish-water wetland of the Gippsland Lakes, south-eastern Australia. Proceedings of the Royal Society of Victoria 120, 403–418.

Boon, P. I., Allen, T., Carr, G., Frood, D., Harty, C., McMahon, A., Mathews, S., Rosengren, N., Sinclair, S., White, M., and Yugovic, J. (2015). Coastal wetlands of Victoria, south-eastern Australia: providing the inventory and condition information needed for their effective management and conservation. Aquatic Conservation 25, 454–479.
Coastal wetlands of Victoria, south-eastern Australia: providing the inventory and condition information needed for their effective management and conservation.Crossref | GoogleScholarGoogle Scholar |

Boon, P. I., Cook, P., and Woodland, R. (2016). The challenges posed by chronic environmental change in the Gippsland Lakes Ramsar site. Marine and Freshwater Research 67, 721–737.
The challenges posed by chronic environmental change in the Gippsland Lakes Ramsar site.Crossref | GoogleScholarGoogle Scholar |

Brinson, M. M., and Rheinhardt, R. (1996). The role of reference wetlands in functional assessment and mitigation. Ecological Applications 6, 69–76.
The role of reference wetlands in functional assessment and mitigation.Crossref | GoogleScholarGoogle Scholar |

Burdick, D., and Konisky, R. A. (2003). Determinants of expansion for Phragmites australis, common reed, in natural and impacted coastal marshes. Estuaries 26, 407–416.
Determinants of expansion for Phragmites australis, common reed, in natural and impacted coastal marshes.Crossref | GoogleScholarGoogle Scholar |

Carter, E. S., White, S. M., and Wilson, A. M. (2008). Variation in groundwater salinity in a tidal salt marsh basin, North Inlet estuary, South Carolina. Estuarine, Coastal and Shelf Science 76, 543–552.
Variation in groundwater salinity in a tidal salt marsh basin, North Inlet estuary, South Carolina.Crossref | GoogleScholarGoogle Scholar |

Carus, J., Heuner, M., Paul, M., and Schröder, B. (2017). Plant distribution and stand characteristics in brackish marshes: unravelling the roles of abiotic factors and interspecific competition. Estuarine, Coastal and Shelf Science 196, 237–247.
Plant distribution and stand characteristics in brackish marshes: unravelling the roles of abiotic factors and interspecific competition.Crossref | GoogleScholarGoogle Scholar |

Chen, Q., Wang, Y. D., Zou, C. B., and Wang, Z. L. (2017). Aboveground biomass invariance masks significant belowground productivity changes in response to salinization and nitrogen loading in reed marshes. Wetlands 37, 985–995.
Aboveground biomass invariance masks significant belowground productivity changes in response to salinization and nitrogen loading in reed marshes.Crossref | GoogleScholarGoogle Scholar |

Clucas, R. D., and Ladiges, P. Y. (1980). Dieback of Phragmites australis (Common Reed) and increased salinity in the Gippsland Lakes. Publication 292, Ministry for Conservation, Melbourne, Vic., Australia.

Coops, H., Vulink, J. T., and van Nes, E. H. (2004). Managed water levels and the expansion of emergent vegetation along a lakeshore. Limnologica 34, 57–64.
Managed water levels and the expansion of emergent vegetation along a lakeshore.Crossref | GoogleScholarGoogle Scholar |

Crowe, A. S., and Shikaze, S. G. (2004). Linkages between groundwater and coastal wetlands of the Laurentian Great Lakes. Aquatic Ecosystem Health & Management 7, 199–213.
Linkages between groundwater and coastal wetlands of the Laurentian Great Lakes.Crossref | GoogleScholarGoogle Scholar |

Department of Sustainability and Environment (2006). ‘The Index of Wetland Condition Review of Wetland Assessment Methods.’ (Department of Sustainability and Environment: Melbourne, Vic., Australia.)

Ducker, S. C., Brown, V. B., and Calder, D. M. (1977). ‘An Identification of the Aquatic Vegetation in the Gippsland Lakes.’ (School of Botany, The University of Melbourne: Melbourne, Vic., Australia.)

Eller, F., Skalova, H., Caplan, J. S., Bhattarai, G. P., Burger, M. K., Cronin, J. T., Guo, W. Y., Guo, X., Hazelton, E. L. G., and Kettering, K. M. (2017). Cosmopolitan species as models for ecophysiological responses to global change: the common reed Phragmites australis. Frontiers in Plant Science 8, 1833.
Cosmopolitan species as models for ecophysiological responses to global change: the common reed Phragmites australis.Crossref | GoogleScholarGoogle Scholar |

Ellis, J. and Lee, T. (2002). ‘Casting the Net: Early Fishing Families of the Gippsland Coast.’ (Lakes Entrance Family History Resource Centre: Lakes Entrance, Vic., Australia.)

Engloner, A. I. (2009). Structure, growth dynamics and biomass of reed (Phragmites australis) – a review. Flora 204, 331–346.
Structure, growth dynamics and biomass of reed (Phragmites australis) – a review.Crossref | GoogleScholarGoogle Scholar |

Environment Protection Authority Victoria (2008). Gippsland Lakes Intensive Water Quality Monitoring Program 2006–07. Publication 1241, EPA Victoria, Melbourne, Vic., Australia.

Environment Protection Authority Victoria (2015). ‘Gippsland Lakes and Catchment Literature Review.’ (EPA Victoria: Melbourne, Vic., Australia.)

Fogli, S., Marchesini, R., and Gerdol, R. (2002). Reed (Phragmites australis) decline in a brackish wetland in Italy. Marine Environmental Research 53, 465–479.
Reed (Phragmites australis) decline in a brackish wetland in Italy.Crossref | GoogleScholarGoogle Scholar |

Fryer, J. (1973). Development of the Gippsland Lakes entrance since 1851. Proceedings of the Royal Society of Victoria 85, 125–133.

Ganf, G., White, S., and Oliver, R. (2010). Allocating water to the wetlands of the Murray Valley to maximise aquatic plant species diversity. In ‘Ecosystem Response Modelling in the Murray–Darling Basin’. (Eds N. Saintilan and I. Overton.) pp. 279–299. (CSIRO Publishing: Melbourne, Vic., Australia.)

Gedan, K. B., Kirwan, M. L., Wolanski, E., Barbier, E. B., and Silliman, B. R. (2011). The present and future role of coastal wetland vegetation in protecting shorelines: answering recent challenges to the paradigm. Climatic Change 106, 7–29.
The present and future role of coastal wetland vegetation in protecting shorelines: answering recent challenges to the paradigm.Crossref | GoogleScholarGoogle Scholar |

Gippsland Ports (2018). Tide predictions. Available at https://www.gippslandports.vic.gov.au/boating/waves-tides-and-weather/tide-predictions/ [Verified 11 June 2018].

Gorai, M., Vadel, A., Neffati, M., and Khemira, H. (2007). The effect of sodium chloride salinity on the growth, water status and ion content of Phragmites communis Trin. Pakistan Journal of Biological Sciences 10, 2225–2230.
The effect of sodium chloride salinity on the growth, water status and ion content of Phragmites communis Trin.Crossref | GoogleScholarGoogle Scholar |

Gorai, M., Ehnajeh, M., Khemira, H., and Neffati, M. (2011). Influence of NaCl-salinity on growth, water relations and solute accumulation in Phragmites australis. Acta Physiologiae Plantarum 33, 963–971.
Influence of NaCl-salinity on growth, water relations and solute accumulation in Phragmites australis.Crossref | GoogleScholarGoogle Scholar |

Greenwood, M. E., and MacFarlane, G. R. (2006). Effects of salinity and temperature on the germination of Phragmites australis, Juncus kraussii, and Juncus acutus: implications for estuarine restoration initiatives. Wetlands 26, 854–861.
Effects of salinity and temperature on the germination of Phragmites australis, Juncus kraussii, and Juncus acutus: implications for estuarine restoration initiatives.Crossref | GoogleScholarGoogle Scholar |

Gregory, J. W. (1903). ‘The Geography of Victoria.’ (Whitcombe & Tombs: Melbourne, Vic., Australia.)

Guan, B., Yu, J. B., Hou, A. X., Han, G. X., Wang, G. M., Qu, F. Z., Xia, J. B., and Wang, X. H. (2017). The ecological adaptability of Phragmites australis to interactive effects of water level and salt stress in the Yellow River delta. Aquatic Ecology 51, 107–116.
The ecological adaptability of Phragmites australis to interactive effects of water level and salt stress in the Yellow River delta.Crossref | GoogleScholarGoogle Scholar |

Harris, G. P. (2001). Biogeochemistry of nitrogen and phosphorus in Australian catchments, rivers and estuaries: effects of land use and flow regulation and comparisons with global patterns. Marine and Freshwater Research 52, 139–149.
Biogeochemistry of nitrogen and phosphorus in Australian catchments, rivers and estuaries: effects of land use and flow regulation and comparisons with global patterns.Crossref | GoogleScholarGoogle Scholar |

Hart, T. S. (1921). The Gippsland Lakes country: physiographical features. Victorian Naturalist 38, 75–82.

Hawkins, R. (2000). ‘Creeks and Harbours of the Gippsland Lakes and Eastern Gippsland’, 2nd edn. (In-depth Publications: Metung, Vic., Australia.)

Hellings, S. E., and Gallagher, J. L. (1992). The effects of salinity and flooding on Phragmites australis. Journal of Applied Ecology 29, 41–49.
The effects of salinity and flooding on Phragmites australis.Crossref | GoogleScholarGoogle Scholar |

Herbert, E. R., Boon, P. I., Burgin, A. J., Neubauer, S. C., Franklin, R. B., Ardón, M., Hopfensperger, K. N., Lamers, L. P. M., and Gell, P. (2015). A global perspective on wetland salinization: ecological consequences of a growing threat to freshwater wetlands. Ecosphere 6, art206.
A global perspective on wetland salinization: ecological consequences of a growing threat to freshwater wetlands.Crossref | GoogleScholarGoogle Scholar |

Holmes, G., Hall, N. E., Gendall, A. R., Boon, P. I., and James, E. (2016). Using transcriptomics to identify differential gene expression in response to salinity among Australian Phragmites australis clones. Functional Plant Ecology 7, 432.
Using transcriptomics to identify differential gene expression in response to salinity among Australian Phragmites australis clones.Crossref | GoogleScholarGoogle Scholar |

Honnay, O., and Bossuyt, B. (2005). Prolonged clonal growth: escape route or route to extinction? Oikos 108, 427–432.
Prolonged clonal growth: escape route or route to extinction?Crossref | GoogleScholarGoogle Scholar |

Howard, R. J., and Rafferty, P. S. (2006). Clonal variation in response to salinity and flooding stress in four marsh macrophytes of the northern Gulf of Mexico, USA. Environmental and Experimental Botany 56, 301–313.
Clonal variation in response to salinity and flooding stress in four marsh macrophytes of the northern Gulf of Mexico, USA.Crossref | GoogleScholarGoogle Scholar |

Hroudová, Z., Zákravsky, P., and Flegrová, M. (2014). The tolerance to salinity and nutrient supply in four European Bolboschoenus species (B. maritimus, B. laticarpus, B. planiculmis and B. yagara) affects their vulnerability or expansiveness. Aquatic Botany 112, 66–75.
The tolerance to salinity and nutrient supply in four European Bolboschoenus species (B. maritimus, B. laticarpus, B. planiculmis and B. yagara) affects their vulnerability or expansiveness.Crossref | GoogleScholarGoogle Scholar |

Hurlbert, S. H. (1984). Pseudoreplication and the design of ecological field experiments. Ecological Monographs 54, 187–211.
Pseudoreplication and the design of ecological field experiments.Crossref | GoogleScholarGoogle Scholar |

Hurry, C., James, E. A., and Thompson, R. M. (2013). Connectivity, genetic structure and stress response of Phragmites australis: issues for restoration in a salinising wetland system. Aquatic Botany 104, 138–146.
Connectivity, genetic structure and stress response of Phragmites australis: issues for restoration in a salinising wetland system.Crossref | GoogleScholarGoogle Scholar |

James, E. A., Jordan, R., and Griffin, P. C. (2013). Spatial genetic analysis of two polyploid macrophytes reveals high connectivity in a modified wetland. Freshwater Biology 58, 2102–2113.
Spatial genetic analysis of two polyploid macrophytes reveals high connectivity in a modified wetland.Crossref | GoogleScholarGoogle Scholar |

King, G. M. (1981). Tidal scour in the Gippsland Lakes. Proceedings of the Royal Society of Victoria 92, 131–136.

Li, S.-H., Ge, Z.-M., Xie, L.-N., Yuan, L., Wang, D.-Q., Li, X.-Z., and Zhang, L.-Q. (2018). Ecophysiological response of native and exotic salt marsh vegetation to waterlogging and salinity: implications for the effects of sea-level rise. Nature Scientific Reports 8, 2441.
Ecophysiological response of native and exotic salt marsh vegetation to waterlogging and salinity: implications for the effects of sea-level rise.Crossref | GoogleScholarGoogle Scholar |

Lillebø, A. I., Pardal, M. A., Neto, J. M., and Marques, J. C. (2003). Salinity as the major factor affecting Scirpus maritimus annual dynamics: evidence from field data and greenhouse experiment. Aquatic Botany 77, 111–120.
Salinity as the major factor affecting Scirpus maritimus annual dynamics: evidence from field data and greenhouse experiment.Crossref | GoogleScholarGoogle Scholar |

Lissner, J., and Schierup, H. H. (1997). Effects of salinity on the growth of Phragmites australis. Aquatic Botany 55, 247–260.
Effects of salinity on the growth of Phragmites australis.Crossref | GoogleScholarGoogle Scholar |

Lissner, J., Schierup, H. H., Comin, F. A., and Astorga, V. (1999). Effect of climate on the salt tolerance of two Phragmites australis populations. I. Growth, inorganic solutes, nitrogen relations and osmoregulation. Aquatic Botany 64, 317–333.
Effect of climate on the salt tolerance of two Phragmites australis populations. I. Growth, inorganic solutes, nitrogen relations and osmoregulation.Crossref | GoogleScholarGoogle Scholar |

Liu, Q., and Mou, X. (2016). Interactions between surface water and groundwater: key processes in ecological restoration of degraded coastal wetlands caused by reclamation. Wetlands 36, 95–102.
Interactions between surface water and groundwater: key processes in ecological restoration of degraded coastal wetlands caused by reclamation.Crossref | GoogleScholarGoogle Scholar |

Long, A. L., Kettenring, K. M., Hawkins, C. P., and Neale, C. M. U. (2017). Distribution and drivers of a widespread, invasive wetland grass, Phragmites australis, in wetlands of the Great Salt Lake, Utah, USA. Wetlands 37, 45–57.
Distribution and drivers of a widespread, invasive wetland grass, Phragmites australis, in wetlands of the Great Salt Lake, Utah, USA.Crossref | GoogleScholarGoogle Scholar |

Lynch, E. A., and Saltonstall, K. (2002). Paleoecological and genetic analyses provide evidence for recent colonization of native Phragmites australis populations in a Lake Superior wetland. Wetlands 22, 637–646.
Paleoecological and genetic analyses provide evidence for recent colonization of native Phragmites australis populations in a Lake Superior wetland.Crossref | GoogleScholarGoogle Scholar |

Matoh, T., Matsushita, N., and Takahasi, E. (1988). Salt tolerance in the reed plant Phragmites communis. Physiologia Plantarum 72, 8–14.
Salt tolerance in the reed plant Phragmites communis.Crossref | GoogleScholarGoogle Scholar |

Meyerson, L. A., Saltonstall, K., and Chambers, R. M. (2009). Phragmites australis in eastern North America: a historical and ecological perspective. In ‘Human Impacts on Salt Marshes: a Global Perspective’. (Eds B. R. Silliman, E. D. Grosholz, and M. D. Bertness.) pp. 57–82. (University of California Press: Berkeley, CA, USA.)

Möller, I., and Spencer, T. (2002). Wave dissipation over macro-tidal saltmarshes: effects of marsh edge typology and vegetation change. Journal of Coastal Research 36, 506–521.
Wave dissipation over macro-tidal saltmarshes: effects of marsh edge typology and vegetation change.Crossref | GoogleScholarGoogle Scholar |

Morris, K., Boon, P. I., Raulings, E. J., and White, S. E. (2008). Floristic shifts in wetlands: the effects of environmental variables on the interaction between Phragmites australis (common reed) and Melaleuca ericifolia (swamp paperbark). Marine and Freshwater Research 59, 187–204.
Floristic shifts in wetlands: the effects of environmental variables on the interaction between Phragmites australis (common reed) and Melaleuca ericifolia (swamp paperbark).Crossref | GoogleScholarGoogle Scholar |

Nada, R. M., Kedr, A. H. A., Serag, M. S., and El-Nagar, N. A. (2015). Growth, photosynthesis and stress-inducible genes of Phragmites australis (Cav.) Trin. ex Steudel from different habitats. Aquatic Botany 124, 54–62.
Growth, photosynthesis and stress-inducible genes of Phragmites australis (Cav.) Trin. ex Steudel from different habitats.Crossref | GoogleScholarGoogle Scholar |

Packer, J. G., Meyerson, L. A., Skálova, H., Pyšek, P., and Kueffer, C. (2017). Biological flora of the British Isles: Phragmites australis. Journal of Ecology 105, 1123–1162.
Biological flora of the British Isles: Phragmites australis.Crossref | GoogleScholarGoogle Scholar |

Phillipson, S., Duncan, L., and Hardie, R. (2014). Mitchell River silt jetties shoreline protection and enhancement project. Report to Parks Victoria, Alluvium, Melbourne, Vic., Australia. Available at http://www.loveourlakes.net.au/wp-content/uploads/2018/06/14029_PV_mitchell_river_silt_jetties_FINAL_v3.1.pdf [Verified 6 September 2018].

Quinn, G. P., and Keough, M. J. (2002). ‘Experimental Design and Data Analysis for Biologists.’ (Cambridge University Press: Cambridge, UK.)

Raulings, E., Morris, K., Roache, M., and Boon, P. I. (2010). The importance of water regimes operating at small spatial scales for the diversity and structure of wetland vegetation. Freshwater Biology 55, 701–715.
The importance of water regimes operating at small spatial scales for the diversity and structure of wetland vegetation.Crossref | GoogleScholarGoogle Scholar |

Raulings, E., Morris, K., Roache, M., and Boon, P. I. (2011). Is hydrological manipulation an effective management tool for rehabilitating chronically flooded, brackish-water wetlands? Freshwater Biology 56, 2347–2369.
Is hydrological manipulation an effective management tool for rehabilitating chronically flooded, brackish-water wetlands?Crossref | GoogleScholarGoogle Scholar |

Rawlinson, T. E. (1863). Report on the entrance to the Gippsland Lakes and notes on the coast and lakes of Gippsland. Transactions of the Royal Society of Victoria 6, 84–98.

Rea, N., and Ganf, G. G. (1994). The role of sexual reproduction and water regime in shaping the distribution patterns of clonal emergent aquatic vegetation. Australian Journal of Marine and Freshwater Research 45, 1469–1479.
The role of sexual reproduction and water regime in shaping the distribution patterns of clonal emergent aquatic vegetation.Crossref | GoogleScholarGoogle Scholar |

Reid, M., Fluin, J., Ogden, R., Tibby, J., and Kershaw, P. (2002). Long-term perspectives on human impacts on floodplain–river ecosystems, Murray–Darling Basin, Australia. Verhandlungen des Internationalen Verein Limnologie 28, 1–7.

Rickey, M. A., and Anderson, R. C. (2004). Effects of nitrogen addition on the invasive grass Phragmites australis and a native competitor Spartina pectinata. Journal of Applied Ecology 41, 888–896.
Effects of nitrogen addition on the invasive grass Phragmites australis and a native competitor Spartina pectinata.Crossref | GoogleScholarGoogle Scholar |

Roberts, J. (2000). Changes in Phragmites australis in south-eastern Australia: a habitat assessment. Folia Geobotanica 35, 353–362.
Changes in Phragmites australis in south-eastern Australia: a habitat assessment.Crossref | GoogleScholarGoogle Scholar |

Robinson, R. W., James, E. A., and Boon, P. I. (2012). Population structure in the woody wetland plant Melaleuca ericifolia Sm. (Myrtaceae): an analysis using historical aerial photographs and molecular techniques. Australian Journal of Botany 60, 9–19.
Population structure in the woody wetland plant Melaleuca ericifolia Sm. (Myrtaceae): an analysis using historical aerial photographs and molecular techniques.Crossref | GoogleScholarGoogle Scholar |

Rogers, K. (2011). Vegetation. In ‘Floodplain Wetland Biota in the Murray–Darling Basin: Water and Habitat Requirements’. (Eds K. Rogers and T. J. Ralph.) pp. 17–82. (CSIRO Publishing: Melbourne, Vic., Australia.)

Rosengren, N. (1984). Sites of geological and geomorphological significance in the Gippsland Lakes Catchment. Publication 402, Environmental Studies Series, Ministry for Conservation, Department of Conservation Forests and Lands, Melbourne, Vic., Australia.

Sainty, G. R., and Jacobs, S. W. L. (2003). ‘Waterplants in Australia’, 4th edn. (Sainty and Associates: Sydney, NSW, Australia.)

Saunders, K. M., Hodgson, D. A., Harrison, J., and McMinn, A. (2008). Palaeoecological tools for improving the management of coastal ecosystems: a case study from Lake King (Gippsland Lakes) Australia. Journal of Paleolimnology 40, 33–47.
Palaeoecological tools for improving the management of coastal ecosystems: a case study from Lake King (Gippsland Lakes) Australia.Crossref | GoogleScholarGoogle Scholar |

Schenck, F. R., Hanley, T. C., Beighley, R. E., and Hughes, A. R. (2018). Phenotypic variation among invasive Phragmites australis populations does not influence salinity tolerance. Estuaries and Coasts 41, 896–907.

Shepard, C. C., Crain, C. M., and Beck, M. W. (2011). The protective role of coastal marshes: a systematic review and meta-analysis. PLoS One 6, e27374.
The protective role of coastal marshes: a systematic review and meta-analysis.Crossref | GoogleScholarGoogle Scholar |

Sinclair, S., and Boon, P. I. (2012). Changes in the area of coastal marsh in Victoria since the mid 19th century. Cunninghamia 12, 153–176.

Sjerp, E., Martin, B., Riedel, P., and Bird, E. C. F. (2002). Gippsland Lakes shore erosion and revegetation strategy. Report to Gippsland Coastal Board. (Ethos NRM: Bairnsdale, Vic., Australia.) Available at https://www.google.com/url?sa=t&rct=j&q=&esrc=s&source=web&cd=1&cad=rja&uact=8&ved=2ahUKEwj3n7H2s47dAhXUdt4KHWeCBEwQFjAAegQIARAC&url=http%3A%2F%2Fwww.wellington.vic.gov.au%2Ffiles%2Fbd1fbbb5-40fc-4bb3-84bb-a8e900fb441c%2FGippsland-Lakes-Shore-Erosion-and-Revegetation-Strategy.pdf&usg=AOvVaw2Mh9YmgjhFC6RLBGh8jdmS [Verified 28 August 2018].

Spalding, M. D., McIvor, A. L., Beck, M. W., Koch, E. W., Möller, I., Reed, D. J., Rubinoff, P., Spencer, T., Tolhurst, T. J., Wamsley, T. V., van Wesenbeeck, B. K., Wolanski, E., and Woodroffe, C. D. (2014). Coastal ecosystems: a critical element of risk reduction. Conservation Letters 7, 293–301.
Coastal ecosystems: a critical element of risk reduction.Crossref | GoogleScholarGoogle Scholar |

Sutter, L. A., Chambers, R. M., and Perry, J. E. (2015). Seawater intrusion mediates species transition in low salinity tidal marsh vegetation. Aquatic Botany 122, 32–39.
Seawater intrusion mediates species transition in low salinity tidal marsh vegetation.Crossref | GoogleScholarGoogle Scholar |

Synan, P. (1989). ‘Highways of Water: How Shipping on the Lakes Shaped Gippsland.’ (Landmark Press: Drouin, Vic., Australia.)

Takahashi, R., Nishio, T., Ichizen, N., and Takano, T. (2007). Salt-tolerant reed plants contain lower Na+ and higher K+ than salt-sensitive reed plants. Acta Physiologiae Plantarum 29, 431–438.
Salt-tolerant reed plants contain lower Na+ and higher K+ than salt-sensitive reed plants.Crossref | GoogleScholarGoogle Scholar |

Thoms, M. C., Ogden, R. W., and Reid, M. A. (1999). Establishing the condition of lowland floodplain rivers: a palaeoecological approach. Freshwater Biology 41, 407–423.
Establishing the condition of lowland floodplain rivers: a palaeoecological approach.Crossref | GoogleScholarGoogle Scholar |

Underwood, A. J. (1981). Techniques of analysis of variance in experimental marine biology and ecology. Oceanography and Marine Biology – an Annual Review 19, 513–605.

United States Environmental Protection Agency (2006). Salinity. In ‘Volunteer Estuary Monitoring Manual. A Methods Manual’, 2nd edn. Available at https://www.epa.gov/sites/production/files/2015-09/documents/2009_03_13_estuaries_monitor_chap14.pdf [Verified 1 June 2018].

van der Putten, W. H. (1997). Die-back of Phragmites australis in European wetlands: an overview of the European Research Programme on Reed Die-Back and Progression (1993–1994). Aquatic Botany 59, 263–275.
Die-back of Phragmites australis in European wetlands: an overview of the European Research Programme on Reed Die-Back and Progression (1993–1994).Crossref | GoogleScholarGoogle Scholar |

Vasquez, E. A., Glenn, E. P., Brown, J. J., Guntenspergen, G. R., and Nelson, S. G. (2005). Salt tolerance underlies the cryptic invasion of North American salt marshes by an introduced haplotype of the common reed Phragmites australis (Poaceae). Marine Ecology Progress Series 298, 1–8.
Salt tolerance underlies the cryptic invasion of North American salt marshes by an introduced haplotype of the common reed Phragmites australis (Poaceae).Crossref | GoogleScholarGoogle Scholar |

Vasquez, E. A., Glenn, E. P., Guntenspergen, G. R., Brown, J. J., and Nelson, S. G. (2006). Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient. American Journal of Botany 93, 1784–1790.
Salt tolerance and osmotic adjustment of Spartina alterniflora (Poaceae) and the invasive M haplotype of Phragmites australis (Poaceae) along a salinity gradient.Crossref | GoogleScholarGoogle Scholar |

Webster, I. T., Parslow, J. S., Grayson, R. B., Molloy, R. P., Andrewartha, J., Sakov, P., Tan, K. S., Walker, S. J., and Wallace, B. B. (2001). Gippsland Lakes environmental study: assessing options for improving water quality and ecological function. Report to the Gippsland Coastal Board, Bairnsdale. (CSIRO: Adelaide, SA, Australia.) Available at http://www.monitor2manage.com.au/userdata/downloads/p_/Gipps26Nov01.pdf [Verified 28 August 2018].

Wells, J. (1986). ‘Gippsland: A People, A Place and Their Past.’ (Landmark Press: Drouin, Vic., Australia.)

Werner, A. D., and Lockington, D. A. (2006). Tidal impacts on riparian salinities near estuaries. Journal of Hydrology 328, 511–522.
Tidal impacts on riparian salinities near estuaries.Crossref | GoogleScholarGoogle Scholar |

Willby, N. J. (2011). From metrics to Monet: the need for an ecologically meaningful guiding image. Aquatic Conservation 21, 601–603.
From metrics to Monet: the need for an ecologically meaningful guiding image.Crossref | GoogleScholarGoogle Scholar |

Willis, J. H. (1970). ‘A Handbook of Plants in Victoria. Volume 1: Ferns, Conifers and Monocotyledons’, 2nd edn. (The University of Melbourne Press: Melbourne, Vic., Australia.)

Xiao, K., Li, H., Wilson, A. M., Xia, Y., Wan, L., Zheng, C., Ma, Q., Wang, C., Wang, X., and Jiang, X. (2017). Tidal groundwater flow and its ecological effects in a brackish marsh at the mouth of a large sub-tropical river. Journal of Hydrology 555, 198–212.
Tidal groundwater flow and its ecological effects in a brackish marsh at the mouth of a large sub-tropical river.Crossref | GoogleScholarGoogle Scholar |

Yang, Z. F., Xie, T., and Liu, Q. (2014). Physiological responses of Phragmites australis to the combined effects of water and salinity stress. Ecohydrology 7, 420–426.
Physiological responses of Phragmites australis to the combined effects of water and salinity stress.Crossref | GoogleScholarGoogle Scholar |

Zar, J. D. (1984). ‘Biostatistical Analysis’, 2nd edn. (Prentice Hall: Englewood Cliffs, NJ, USA.)

Zhen, W. B., and Ma, Q. H. (2009). Proline metabolism in response to salt stress in common reed [Phragmites australis (Cav.) Trin. ex Steud]. Botanica Marina 52, 341–347.
Proline metabolism in response to salt stress in common reed [Phragmites australis (Cav.) Trin. ex Steud].Crossref | GoogleScholarGoogle Scholar |