1 The construction of a raised dike across the coastal saltmarshes of Odiel (Huelva, south-west Spain) in 1977 divided a uniform area of low-lying sediment into two lagoons with very different drainage properties. To the east of the dike, there is rapid drainage into the estuarine channel of the Odiel and Tinto rivers, whereas to the west, the development of sand spits has impeded drainage such that standing water persists long after high tides, despite the slightly higher elevation of this area. 2 Both lagoons have been colonized by isolated clones of Spartina maritima, which have locally enhanced accretion to form domed tussocks (`nucleation'). The diameter of the tussocks was highly correlated with the elevation of their sediment surface, the larger tussocks having been the earlier colonists in a generally accreting system. Spartina in the interior of the tussocks showed reduced tiller density and vigour. 3 Only in the better-drained lagoon have the central, higher areas of the Spartina tussocks been invaded by Arthrocnemum perenne, although seed is freely available in both lagoons. Arthrocnemum formed a sprawling, dense canopy and a superficial, relatively impenetrable root system; it rapidly suppressed the remaining tillers of Spartina, eventually leaving only a fringe around the edge of the tussock. Areas invaded by Arthrocnemum were characterized by a superficial layer (10 cm) of oxidizing sediment $(E_h > + 80 mV)$. Spartina-dominated areas in both lagoons remained highly reducing, even in the surface layers ($E_h < - 150 mV$). 4 In terms of successional mechanisms, primary colonization by Spartina maritima facilitates the invasion by Arthrocnemum, which only becomes established from seed on relatively well-drained, oxidising sediments. The interaction is, however, more complicated than this: declining tiller density within the tussocks prior to invasion by Arthrocnemum is consistent with an inhibition mechanism, and the superior competitive ability of Arthrocnemum, once established, suggests that a tolerance mechanism may also operate

1 Salt marshes in south-west Spain are being invaded by Spartina densi¯ora, a South American introduction, although the native S. maritima still dominates many lower marshes. A transplant experiment was used to investigate the means by which physical and chemical factors may determine lower vegetation limits in the tidal frame. Both species were transplanted from a mid-level marsh to lower, unvegetated tidal¯ats. 2 The survival and growth of transplanted clumps and their constituent tillers were monitored on an elevational gradient. The photosynthetic competence of transplants was assessed by measurements of leaf gas exchange and the fast kinetics of chlorophyll¯uorescence. Submergence period, salinity, redox potential and sulphide concentration in the sediment were also examined at the transplant sites. 3 Neither species survived for a year at the lowest transplant point ( 1.04 m relative to Spanish Hydrographic Zero). At 1.41 m elevation, S. maritima survived well but all clumps of S. densi¯ora died. At higher elevations ( 1.46±1.67 m), clumps of both species had high survival rates. Tiller growth rates in surviving clumps of both species increased with elevation, but that of S. densi¯ora was more sensitive to low elevation. 4 S. maritima showed no impairment of photosynthetic performance, even at the lowest elevation; its rates of gas exchange were independent of elevation, as were its chlorophyll¯uorescence parameters. In contrast, in S. densi¯ora the rate of CO 2 uptake declined and stress to photosystem II (Fv/Fp) increased at lower elevations; both of these photosynthetic measurements were linearly related to sediment redox potential. Stomatal conductance did not vary with elevation. 5 S. maritima has a potentially wide elevational tolerance and an absolute lower limit substantially below that of S. densi¯ora. The progressively reduced growth and survival of S. densi¯ora at lower levels appear to result primarily from impairment of photosynthesis when root conditions are hypoxic. This eect is mediated through eects on the photochemical apparatus rather than on stomatal resistance to CO 2 uptake. 6 These studies de®ne lower limits to the fundamental niches of the two species in a Mediterranean-type salt marsh and thus provide a basis for future investigations of interactions between them. The correlation between chlorophyll¯uorescence and long-term survival of S. densi¯ora suggests that this short-term measure may be a valuable tool.

The growth responses of the hygro-halophyte A. portulacoides to salinity appear largely to depend on changes in its rate of photosynthetic gas exchange. Photosynthesis appears to be limited mainly through stomatal conductance and hence intercellular CO(2) concentration, rather than by effects on PSII; moderate salinity might stimulate carboxylation capacity. This is in contrast to more extreme halophytes, for which an ability to maintain leaf area can partially offset declining rates of carbon assimilation at high salinity.

Halophytes that are capable of tolerating a wide range of salinity may grow best at intermediate salinities, but the physiological mechanisms underlying positive growth responses to salinity are not clear. This work investigated the growth of Arthrocnemum macrostachyum (Moric) C. Koch (a halophytic C3 shrub) over a wide range of salinities, and the extent to which its responses can be explained by photosynthetic physiology. Growth, gas exchange and chlorophyll fluorescence characteristics of plants were examined in a glasshouse experiment; tissue concentrations of photosynthetic pigments, ash, sodium, potassium, calcium and nitrogen were also determined. Plants showed marked stimulation of growth by salt, with a broad optimum of 171-510 mm NaCl for relative growth rate (RGR). Stimulation of RGR appeared to depend mainly on an increase in specific shoot area, whereas reduced RGR at high salinity (1030 mm) could be attributed to a combination of lower unit shoot (leaf) rate and lower shoot mass fraction. The non-saline treatment plants had the greatest fraction of non-photosynthetic, atrophied surface area. However, net photosynthesis (A) was also stimulated by NaCl, with an optimum of c. 510 mm NaCl. The responses of A to salinity could be accounted for largely by limitation by stomatal conductance (Gs) and intercellular CO(2) concentration (Ci). Even the most hypersaline treatment apparently had no effect on photosystem II (PSII) function, and this resistance could be an important strategy for this halophyte in saline soils. In contrast, Fv/Fm indicated that absence of salt represents an environmental stress for A. macrostachyum and this could be a contributory factor to salt stimulation of A. Notwithstanding the importance of the ability to develop and maintain assimilatory surface area under saline conditions, stimulatory effects on A also appear to be part of a suite of halophytic adaptations in this plant.

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