Jesús M. Castillo

Short-term responses to salinity of an invasive cordgrass

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.

Effects of Salinity on Germination and Seedling Establishment of Endangered Limonium emarginatum (Willd.) O. Kuntze

Abstract The germination and subsequent seedling establishment of Limonium emarginatum, an endangered and endemic halophyte of the Strait of Gibraltar, was studied under exposure to different NaCl concentrations (0, 2, 4, and 6%) in a laboratory experiment. We assessed final germination percentage, number of days to first and final germination, mean time to germinate (MTG), as well as seed viability and seedling survival. Increasing salinity delayed the beginning and ending of germination and reduced final germination percentage, inhibiting germination completely above 2% salinity. L. emarginatum exhibited the greatest germination in fresh water. When seeds were removed from all saline solutions, between 60% and 70% of final germination was recorded, although at hypersalinity, germination viability diminished. Salinity pretreatments had a stimulatory effect on germination since germination speed was higher for the recovery experiment than for the seed germination experiment. Transition between germination and seedling establishment was a critical phase, given that less than 50% of seedlings of L. emarginatum survived in distilled water and 5% survived at 2% salinity.

Nucleation and facilitation in salt pans in Mediterranean salt marshes

Arthrocnemum macrostachyum is a perennial species acting as a primary colonizer of salt pans in Mediterranean high salt marshes. Salicornia ramosissima, an annual, occurs in salt pans under Arthrocnemum canopies and in open areas. The aim of this study was to analyse, in wild populations and a transplant experiment, how S. ramosissima population dy- namics and growth are affected by A. macrostachyum. The environmental conditions within the patches of Arthrocnemum were less stressful than in the open areas, with lower radiation levels and salinity concentrations. In the inner areas of A. macrostachyum patches, density-dependent mortality proc- esses of S. ramosissima seedlings led to low densities of adult individuals with greater morphological development and re- productive success than in open areas. However, at the edges of Arthrocnemum patches facilitation of seedling survival favoured high densities. Environmental stress hindered devel- opment, decreased reproduction and premature death. These results are in agreement with the general theory of factors controlling vegetation distribution that biotic interactions domi- nate in low stress environments, while abiotic interactions dominate under harsher environmental conditions. A. macro- stachyum plays an essential role in the succession in these salt pans, facilitating seed production and stimulating nucleation processes in S. ramosissima.

CLONAL GROWTH AND TILLER DEMOGRAPHY OF THE INVADER CORDGRASS SPARTINA DENSIFLORA BRONGN. AT TWO CONTRASTING HABITATS IN SW EUROPEAN SALT MARSHES

Salt marshes offer a valuable opportunity to study the effects of environment on the clonal growth of species able to live throughout a wide vertical range on the tidal frame. Spartina densiflora is a species native to South America that is invading marshes in SW Europe, NW Africa, and SW North America, where its populations are found from low to high topographic elevations, altering the composition of plant communities. The aim of this study was to increase our knowledge regarding the competitive ability of S. densiflora, analyzing its clonal growth and its ramet demography during 25 months in two expanding populations at a low and a high marsh. Four clonal characteristics were recorded in both populations: (1) dense occupation of available space inside tussocks; (2) lack of a dormant period and high tiller production rates and growth; (3) evidence of physiological integration between ramets; and (4) high rates of sexual reproduction. Furthermore, S. densiflora developed different strategies of clonal growth in contrasting habitats. Tussocks in the low marsh had lower tiller longevity and higher tiller density, natality, and mortality rates, showing faster ramet turnover than in the high marsh. These clonal growth traits would facilitate S. densiflora persistence under the effects of catastrophic events. Spartina densiflora clonal growth traits indicate strong adaptability to different environmental conditions and strong competitive ability, since its tiller dynamics enable it to invade occupied space, and it effectively colonizes new safe-sites by sexual reproduction. These observations help us to understand how this invader has become the most abundant plant in many estuaries of the SW Iberian Peninsula.

Ecophysiology of tidal and non-tidal populations of the invading cordgrass Spartina densiflora: seasonal and diurnal patterns in a Mediterranean climate

Abstract The invasion of salt marshes by alien species poses interesting questions concerning the mechanisms that determine their distribution. Spartina densiflora is one of the species that is invading the Atlantic marshes of SW Iberian Peninsula, where it tolerates a broad range of environmental conditions and it competes with the indigenous vegetation. The high ecological versatility of S. densiflora provides a good tool to study the ecophysiological responses of marsh plants. This article aims to compare the ecophysiological responses of S. densiflora in populations from two contrasting habitats: tidal (middle marsh) and non-tidal (drainage intercepted marsh). The higher values in A (net photosynthesis rate), A / C i (apparent carboxylation efficiency), water use efficiency (intrinsic WUE) and F V / F P (potential photochemical efficiency) recorded at the non-tidal population, except in summertime, agree with previous studies that found better physiological state at lower salinity and higher redox potential. A clear reduction of A at higher irradiance was recorded in spring and summer in the tidal population, coinciding with the lowest ψ (leaf water potential) values. These results help to explain the high primary productivity recorded in S. densiflora populations at brackish marshes in Mediterranean areas. Different responses in gas exchange and chlorophyll fluorescence during summer were recorded for each population. The tidal population showed the maximum values of A in summer. In contrast, the non-tidal population suffered reductions in A , A / C i and F V / F P during summer, when salinity was higher. Thus, salinity limits carbon fixation in S. densiflora non-tidal populations during Mediterranean summer drought. In tidal populations, photosynthesis seems to be more influenced by anoxic conditions. High levels of photoinhibition and low A were recorded on the coldest and less cloudy day, which provoked permanent damages to the photosynthetic apparatus of S. densiflora . This may limit its winter production, as well as its invasion of marshes at higher latitudes in Europe. Finally, discussion about the possible effects of Global Climatic Change on S. densiflora invasion is undertaken.

Contrasting strategies to cope with drought by invasive and endemic species of Lantana in Galapagos

This study compares how Lantana camara, an invasive species, and L. peduncularis, an autochthonous one, cope with drought in Galapagos. Soil surface temperature was the abiotic environmental parameter that best explained variations in photosynthetic stress. Higher soil surface temperatures were recorded in the lowlands and in rain-shadow areas, which were also the driest areas. L. peduncularis, with a shallow root system, behaved as a drought-tolerant species, showing lower relative growth rates, which decreased with leaf water content and higher photosynthetic stress levels in the lowlands and in a northwest rain-shadow area in comparison with higher and wetter locations. Its basal and maximal fluorescences decreased at lower altitudes, reflecting the recorded drops in chlorophyll concentration. In contrast, L. camara with a deep root system behaved as a drought-avoiding species, showing leaf and relative water contents higher than 55% and avoiding permanent damage to its photosynthetic apparatus even in the driest area where it showed very low chlorophyll content. Its relative growth rate decreased more in dry areas in comparison to wetter zones than did that of L. peduncularis, even though it had greater water content. Furthermore, L. camara showed higher water contents, growth rate, and lower photosynthetic stress levels than L. peduncularis in the arid lowlands. Thus, L. peduncularis maintained lower maximum quantum efficiency of photosystem II photochemistry (Fv/Fm) than L. camara even at sunrise, due to higher basal fluorescence values with similar maximal fluorescence, which indicated permanent damage to PSII reaction centres. Our results help to explain the success and limitations of L. camara in the invasion of arid and sub-arid environments.

Phenotypic plasticity of invasive Spartina densiflora (Poaceae) along a broad latitudinal gradient on the Pacific Coast of North America

PREMISE OF THE STUDY Phenotypic acclimation of individual plants and genetic differentiation by natural selection within invasive populations are two potential mechanisms that may confer fitness advantages and allow plants to cope with environmental variation. The invasion of Spartina densiflora across a wide latitudinal gradient from California (USA) to British Columbia (Canada) provides a natural model system to study the potential mechanisms underlying the response of invasive populations to substantial variation in climate and other environmental variables. METHODS We examined morphological and physiological leaf traits of Spartina densiflora plants in populations from invaded estuarine sites across broad latitudinal and climate gradients along the Pacific west coast of North America and in favorable conditions in a common garden experiment. KEY RESULTS Our results show that key foliar traits varied widely among populations. Most foliar traits measured in the field were lower than would be expected under ideal growing conditions. Photosynthetic pigment concentrations at higher latitudes were lower than those observed at lower latitudes. Greater leaf rolling, reduced leaf lengths, and lower chlorophyll and higher carbon concentrations were observed with anoxic sediments. Lower chlorophyll to carotenoids ratios and reduced nitrogen concentrations were correlated with sediment salinity. Our results suggest that the variations of foliar traits recorded in the field are a plastic phenotypic response that was not sustained under common garden conditions. CONCLUSIONS SPARTINA DENSIFLORA shows wide differences in its foliar traits in response to environmental heterogeneity in salt marshes, which appears to be the result of phenotypic plasticity rather than genetic differentiation.

Environmental determination of shoot height in populations of the cordgrass Spartina maritima.

Spartina species tend to exhibit a range of phenotypes, often with short and tall growth forms. Such differences have been attributed variously to environmentally induced phenotypic plasticity and genetic diferentiation between populations. This work examines the basis of height variation inSpartina maritima (Curtis) Fernald at Odiel salt marshes, southwest Spain. Populations from sites with lower sediment redox potentials tended to have significantly taller shoots. Thirty-four natural populations with an 8-fold range of shoot height were transplanted to a common environment on an unvegetated, intertidal plain and shoot height was measured annually for 3 yr. There was a striking convergence in height across populations after transplantation and the change in height in each year of a population was linearly related to its initial height. Most populations grew taller after transplantation, suggesting environmental limitation in their natural habitats. Populations that were originally tall tended to become shorter. The change in shoot height was negatively related to the difference in surface sediment redox potential between their natural sites and the common transplant site. Hypoxic sediments may stimulate stem growth, resulting in improved photosynthetic gas exchange and internal aeration of roots and rhizomes. Although height variation inS. maritima appears mainly to be a result of phenotypic plasticity, a genetic component cannot be ruled out. This study emphasizes the importance of long-term studies, preferably longer than turnover time of shoot populations. The highly plastic growth form ofS. maritima allows it to colonize a wide range of habitats in environmentally heterogeneous salt marshes.

Causes and consequences of salt-marsh erosion in an Atlantic estuary in SW Spain

This study reports on the quantification of horizontal erosion by undermining of slopes in the atlantic mesotidal salt marshes of Odiel, SW Spain, and analyses its causes and consequences. Horizontal erosion has produced considerable losses of salt marsh area, including zones of mature salt marsh. Human pressure, such as from water-borne traffic or the exploitation of the slopes for the capture of bait, increases the natural erosion processes. The role of vegetation in protecting the slopes against erosion is studied. Channel banks covered with plants, many of which belong to species with long-living, above-ground creeping stems, were less eroded than those without vegetation cover. The enormous volume of sediments moved (ca. 7000m3 in one year) could contribute to the silting-up of the navigable channels of the estuary, so that continual dredging is necessary to allow access to shipping. These sediments are highly contaminated, and dredging exposes them more directly to the trophic network of the estuary. There is a considerable loss of natural resources. Finally, the integrated management of this coastal ecosystem is discussed.

Potential of Spartina maritima in Restored Salt Marshes for Phytoremediation of Metals in a Highly Polluted Estuary

Sedimentary abiotic environment, and concentration and stock of nine metals were analyzed in vegetation and sediments to evaluate the phytoremediation capacity of restored Spartina maritima prairies in the highly polluted Odiel Marshes (SW Iberian Peninsula). Samples were collected in two 10 –m long rows parallel to the tidal line at two sediments depths (0–2 cm and 2–20 cm). Metal concentrations were measured by inductively coupled plasma spectroscopy. Iron, aluminum, copper, and zinc were the most concentrated metals. Every metal, except nickel, showed higher concentration in the root zone than at the sediment surface, with values as high as ca. 70 g Fe kg–1. The highest metal concentrations in S. maritima tissues were recorded in its roots (maximum for iron in Spartina roots: 4160.2 ± 945.3 mg kg–1). Concentrations of aluminum and iron in leaves and roots were higher than in superficial sediments. Rhizosediments showed higher concentrations of every metal than plant tissues, except for nickel. Sediment metal stock in the first 20 cm deep was ca. 170.89 t ha–1. Restored S. maritima prairies, with relative cover of 62 ± 6%, accumulated ca. 22 kg metals ha–1. Our results show S. maritima to be an useful biotool for phytoremediation projects in European salt marshes.