Summary 1We examined salt marsh development over 17 years (1984)(1985)(1986)(1987)(1988)(1989)(1990)(1991)(1992)(1993)(1994)(1995)(1996)(1997)(1998)(1999)(2000)(2001) in a rapidly accreting, well-drained lagoon of a macrotidal marsh in south-west Spain. Scattered, elevated tussocks of colonizing Spartina maritima expanded radially until they either coalesced or were separated only by narrow drainage channels. We recorded changes in elevation of the tussocks and investigated the cover and biomass of successive species invading them. 2 Sediment accretion produced a mean annual increase in tussock elevation of 3.5 cm. Sarcocornia perennis had begun to invade the raised centres of the tussocks by 1984, subsequently displacing Spartina maritima radially to become dominant by 1990. A hybrid form of Sarcocornia , which appeared on some of the tussocks occupied by S. perennis in 1997, also expanded radially and had achieved dominance on many of the invaded tussocks by 2001, by virtue of its more erect growth form and rapid accumulation of high above-and below-ground biomass.
3Tussocks not yet invaded by the hybrid remain dominated by S. perennis and have maintained sediment accretion rates and redox potentials similar to those that have been invaded. 4 Genetic analysis, using random amplified DNA (RAPD) markers, indicated that the hybrid was a cross between the diploid Sarcocornia perennis (2 n = 18) and the octaploid S. fruticosa (2 n = 72), a high-marsh species, and suggested that each hybrid individual may have resulted from a separate pollination of an indigenous S. perennis plant. 5 Invasion by the new hybrid thus probably occurred as a result of pollen flow from high-marsh S. fruticosa , some 1 km distant, to the stigmas of the established dominant S. perennis . Succession might therefore be facilitated genetically rather than simply by the enhanced sediment accretion, which ameliorated the effects of submersion and low sediment redox potentials that presumably exclude S. fruticosa from lower parts of the marsh.
Salinity is one of the main chemical factors in salt marshes. Studies focused on the analysis of salinity tolerance of salt marsh plants are very important, since they may help to relate their physiological tolerances with distribution limits in the field. Spartina densiflora is a South America cordgrass, which has started its invasion of the European coastline from the southwestern Iberian Peninsula. In this work, short-term responses in adult tussocks of S. densiflora from southwestern Spain are studied over a wide range of salinity in a greenhouse experiment. Our results point out that S. densiflora has a high tolerance to salinity, showing high growth and net photosynthesis rates from 0.5 to 20 ppt. S. densiflora showed at the lowest salinity (0.5 ppt) high levels of photoinhibition, compensated by higher levels of energy transmission between photosystems. Adaptative mechanisms, as those described previously, would allow it to live in fresh water environments. At the highest salinity (40 ppt), S. densiflora showed a high stress level, reflected in significant decreases in growth, net photosynthesis rate and photochemical efficiency of Photosystem II. These responses support S. densiflora invasion patterns in European estuaries, with low expansion rates along the coastline and faster colonization of brackish marshes and river banks.
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