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.
The South American cordgrass Spartina densiflora is invading European salt marshes getting into contact with the indigenous and endangered lowmarsh dominant, Spartina maritima. This work describes the evolution of the plant zonation during 7 years in a marsh of S. maritima invaded by S. densiflora. S. maritima appeared throughout the whole intertidal gradient from 1.72 to 3.33 m over Spanish Hydrographic Zero (SHZ), showing its higher biomasses and shoot densities at low elevations. In contrast, S. densiflora only invaded upper areas ([+2.59 m SHZ) at the centre of circular tussocks of S. maritima. Above-ground biomass of S. maritima dropped drastically at maximum occupation of space by the alien, and its shoot density and above-ground biomass decreased at S. densiflora zone during the study. The competitive potential of S. densiflora was reflected in high above-and below-ground biomass and shoot densities, accompanied by elevated wrack accumulation and the absence of other marsh plants presented together with S. maritima from areas dominated by S. densiflora. S. densiflora altered the native vegetational zonation pattern through the invasion of the centre of S. maritima tussocks; however, the alien invasion may be limited by the presence of the autochthonous cordgrass at lower elevations.
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