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- Population-level effects of clam harvesting on the seagrass Zostera noltiiPublication . Cabaço, Susana; Alexandre, Ana; Santos, RuiSeagrass declines have been reported worldwide, mostly as a consequence of anthropogenic disturbance. In Ria Formosa lagoon, southern Portugal, the intertidal meadows of Zostera noltii are highly disturbed by clam harvesters. The most common technique used to collect the clams consists of digging and tilling the sediment with a modified knife with a large blade. Here we present both descriptive and experimental evidence of the negative effects of clam harvest on the Z. noltii populations of Ria Formosa. A comparison between disturbed and undisturbed meadows suggests that clam harvesting activities change the species population structure by significantly reducing shoot density and total biomass, particularly during August, when the harvest effort is higher. Experimental harvest revealed a short-term impact on shoot density, which rapidly recovered to control levels during the following month. An experimental manipulation of rhizome fragmentation revealed that plant survival is reduced only when fragmented rhizomes are left with 1 intact internode. Shoot production and rhizome elongation and production of fragmented rhizomes having 2 to 5 internodes were not significantly affected, even though growth and production were lower when only 2 internodes were left. Experimental shoot damage at different positions along the rhizome had a significant effect on plant survival, rhizome elongation, and production only when the apical shoot was removed. Our results show that clam harvest can adversely affect Z. noltii meadows of Ria Formosa while revealing a low modular integration that allows the species to rapidly recover from physical damage.
- Effects of clam harvesting on sexual reproduction of the seagrass Zostera noltiiPublication . Alexandre, Ana; Santos, Rui; Serrão, EsterThe sexual reproduction of Zostera noltii in the Ria Formosa lagoon (southern Portugal) was assessed by quantifying the variation in both flowering effort and evolution of maturation stages during the fertile season. The effects of clam harvesting on the sexual reproduction of Z. noltii were investigated using both descriptive and experimental approaches. Meadows disturbed by clam harvesting were compared with undisturbed meadows in 2 sectors of the lagoon. In both sectors, disturbed meadows showed significantly lower vegetative shoot density and significantly higher reproductive effort. The reproductive effort in the western disturbed meadows was 2-fold higher than in undisturbed meadows, whereas in eastern disturbed meadows it was 4-fold higher. In disturbed meadows, early maturation stages were found until the end of the flowering season and higher seed production was observed. Both the negative effect of clam harvesting on Z. noltii density and the positive effect on its reproductive effort were confirmed by manipulative field experiments. Plots that were experimentally harvested during the fertile season showed significantly lower vegetative shoot density and higher reproductive effort than unharvested plots. Results suggest that Z. noltii responds to clam harvesting disturbance by both increasing its reproductive effort and extending its fertile season.
- Genetic homogeneity in the seagrass Cymodocea nodosa at its northern Atlantic limit revealed through RAPDPublication . Alberto, Filipe; Mata, Leonardo; Santos, RuiRandom amplified polymorphic DNA (RAPD) markers were used to analyse the genetic variability of the dioecious seagrass Cymodocea nodosa Ucria (Ascherson) in the Ria Formosa lagoon, Portugal, the species' northern limit in the Atlantic. Three individuals from each of 6 meadows were genotyped with 28 primers. Meadows described previously as having flower marks were compared with meadows where flowers did not occur. A single polymorphic band, specific for one meadow, was observed in a total of 177 fragments. The lack of genetic variability among meadows both with and without flower indicates that flower production is not associated with a higher level of genetic variation. The genetic homogeneity of C, nodosa in the Ria Formosa suggests a founder effect, produced by a single or a limited number of migrants composing the colonising gene pool, This hypothesis is supported by the geographic isolation from other populations as the nearest populations lie more than 300 km away from the Ria Formosa. The lack of reproductive success of C, nodosa in Ria Formosa natural park and its low genetic variability are important factors in the conservation of this species since recolonisation can only occur through vegetative growth.
- Measuring seagrass photosynthesis: methods and applicationsPublication . Silva, João; Sharon, Yoni; Santos, Rui; Beer, SvenThis review originates from a keynote lecture given at the recent 8th Group for Aquatic Productivity (GAP) workshop held in Eilat, Israel. Here we examine the most important methodologies for photosynthetic measurements in seagrasses and evaluate their applications, advantages and disadvantages, and also point out the most relevant results. The most commonly used methodologies are based on oxygen (O(2)) evolution and chlorophyll fluorescence measurements. O(2)-based methodologies allowed for the first approaches to evaluate seagrass productivity, whereas chlorophyll a fluorescence has more recently become the choice method for in situ experiments, particularly in evaluating photosynthetic responses to light and assessing stress responses. New methodologies have also emerged, such as O(2) optodes, underwater CO(2) flux measurements, geo-acoustic inversion and the eddy correlation technique. However, these new methods still need calibration and validation. Our analysis of the literature also reveals several significant gaps in relevant topics concerning seagrass photosynthesis, namely the complete absence of studies on deep-growing populations that photosynthesise under extreme low light conditions and the uncertainties about the true degree of seagrass carbon limitation, which limits our ability to predict responses to global changes.
- Which demographic traits determine population growth in the invasive brown seaweed Sargassum muticum?Publication . Engelen, Aschwin; Santos, Rui1. Life-history traits commonly associated with plant invasiveness are vegetative reproduction or r-selected traits such as short generation times and high rates of reproduction and individual growth. 2. We used matrix modelling to assess which demographic traits are important for the population growth of an invasive seaweed lacking vegetative reproduction and whether demographic and life-history strategies shift with increased dominance of the invader. The vital rates of one of the most successful invading seaweeds, Sargassum muticum, were investigated monthly for 2 years in intertidal pools dominated by the native brown seaweed Cystoseira humilis and by S. muticum, respectively. In order to speculate about the demographic mechanisms that determine invasiveness of S. muticum, and as the study sites were recently colonized, we assumed that C. humilis and S. muticum pools are proxies for early and late phases of invasion, respectively. 3. Both deterministic and stochastic matrix models showed positive rates of population growth, and rates were significantly higher in the pools dominated by S. muticum than in the ones dominated by C. humilis, indicating demographic changes with invader dominance. The variability of population growth rates and of reproductive and elasticity values of S. muticum was higher in the pools dominated by C. humilis, suggesting invader-driven stabilization of environmental conditions. Generation times of the species increased with invader dominance, supporting invader-stabilized environmental conditions. 4. Elasticity analyses revealed that the most important demographic trait for population growth rate at both levels of invader dominance was the persistence of the non-fertile adult fronds rather than reproduction or growth. No major shifts in the life-history strategy of S. muticum between levels of invader dominance were detected. 5. Synthesis. This study suggests that the invasiveness of S. muticum, a perennial invader without vegetative reproduction, relies on K- rather than r-selected traits and without drastic changes in life-history strategy between phases of invasion.
- Timing and success of reproductive stages in the seagrass Zostera noltiiPublication . Alexandre, Ana; Cabaço, Susana; Santos, Rui; Serrão, EsterThe timing and success of sexual reproduction of the seagrass Zostera noltii was investigated at the Ria Formosa lagoon, Portugal. Thirty plants were tagged and monitored individually through time to determine in situ the development time of each maturation stage, from the emergence of the flowers to the production of seeds. The overall process of flowering and fruiting lasted 47 ± 4 days, during which formation and maturation of the fruits was the most time-consuming stage (27 ± 2 days). Spathe success, i.e. the percentage of spathes that produced seeds, was 22 ± 4% while spathe mortality was 34 ± 6%. A considerable percentage of spathes (37 ± 7%) was lost through leaf detachment, but some of these may still add to the reproductive success of the species by contributing to the species recruitment within the source meadow or elsewhere. Meadow seed production (MSP) of Z. noltii was estimated to be 312 ± 66 seeds m−2, whereas the potential seed production of the studied meadow was 2623 seeds m−2. Under laboratory conditions, 70% of Z. noltii seeds germinated within 26 days, but only 10% reached the seedling stage due to the high mortality of germlings. Fertility, defined as the probability of a seed to originate a new plant, was estimated to be 14 × 10−4, which is higher than what is expected for most seagrasses.
- Daily variation patterns in seagrass photosynthesis along a vertical gradientPublication . Silva, João; Santos, RuiThe relationship between the available photosynthetic active radiation (PAR) and the photosynthetic yield of Photosystem II (PSII), measured by chlorophyll a fluorescence, was assessed in the intertidal seagrasses of Ria Formosa, a coastal lagoon in southern Portugal. Most of the lagoon's intertidal is occupied by a monospecific population of Zostera noltii (Hornemann), which occupies a vertical gradient of about 2 m. The upper distribution limit of this species comprises the edge of a Spartina maritima (Curtis) Fernald saltmarsh and the lower limit the transition to the subtidal, dominated by Cymodocea nodosa (Ucria) Ascherson. Diurnal changes in the pattern of rapid-light curves (RLCs) was investigated with pulse amplitude-modulated (PAM) fluorometry in Z. noltii and in C. nodosa in the upper and lower intertidal. The light reactions of photosynthesis were assessed by fitting photosynthesis-irradiance (P-I) models to the RLCs. The photosynthetic parameters of Z noltii revealed that this plant species exhibits sun- and shade-type responses in its upper and lower vertical distribution limits, respectively. At the lower distribution limit in low light, the initial slopes of all RLCs were significantly higher than at the upper site, but decreased with increasing irradiances, while at the upper distribution limit, although lower, the slopes were unaffected by increasing irradiance. C. nodosa presented a typical shade-type response, as evidenced by the daily variation and light dependence of both photosynthetic efficiency and optimal quantum yield. The relationship between the maximum electron-transport rate and irradiance suggests that this species is strongly light-limited. We suggest that attempts to characterize the photosynthetic behaviour of an intertidal meadow should consider both diurnal fluctuations in the plants' photosynthetic activity as well as its vertical distribution frequency.
- Habitat differences in the timing of reproduction of the invasive alga sargassum muticum (phaeophyta, sargassaceae) over tidal and lunar cyclesPublication . Monteiro, C. A.; Engelen, Aschwin; Serrão, Ester; Santos, RuiSargassum muticum (Yendo) Fensholt is an invasive species that is firmly established on intertidal and subtidal rocky shores of Europe and the Pacific coast of North America. Local success and spread of S. muticum is thought to rely on its reproductive potential that seems dependent on exogenous factors like tidal and lunar cycles. This study is the first to compare the reproductive patterns (periodicity of egg expulsion and embryo settlement) of this invader in two different habitats: the middle and low intertidal. The combination of monthly, daily, and tidal samples at triplicate sites within each habitat showed a semilunar periodicity of egg expulsion and embryo settlement coincident with increasing tidal amplitude just before full and new moons. In both habitats, duration of each egg expulsion event was 1 week, and embryo settlement occurred during the first daily low tide and with the incoming high tide during spring tides. However, both expulsion and settlement started 1–2 d earlier, expulsion saturation was faster, and settlement was higher in the mid- compared to the low intertidal. Our results suggest that the exact timing of gamete expulsion and embryo release of S. muticum responds to local factors, including tidal cues, which result in differences between mid- and lowintertidal habitats.
- Oxidative stress and quantum yield efficiency in the intertidal seagrass zostera noltiiPublication . Silva, João; Barrote, Isabel; Santos, RuiWe investigated the combined effects of several environmental stressors in the photosynthetic performance and in the activation of biochemical defense mechanisms in the intertidal seagrass Zostera noltii in Ria Formosa coastal lagoon (southern Portugal). The maximum (Fv/Fm) and the effective (F’v/F’m) quantum use efficiencies of PSII were sampled monthly in both neap and spring tides over one year. Other fluorescence parameters, such as the Stern-Volmer non-photochemical quenching (NPQ) and the novel parameter LNP (which expresses the general decrease in PSII photochemical activity in the light) were derived from quantum use efficiency measurements. Sampling for antioxidant enzymes activity, pigments, soluble protein and malondialdehyde (MDA) was conducted in parallel. Reactive oxygen species (ROS) are formed as a normal part of the metabolism. An increment on ROS formation is a common response to those stresses and can cause several types of damage, namely lipid peroxidation. MDA is a product of the peroxidation of membrane lipids and thus is commonly used as an indicator of oxidative stress. Carotenoids and antioxidant enzymes such as ascorbate peroxidase (APx) are part of the plants’ antioxidative system. Higher content of carotenoids indicate a higher photoprotection and increased activities of ROS scavenging enzymes such as APx are correlated with stress tolerance. The relationships among critical environmental parameters (irradiance, temperature, air exposure), oxidative stress, antioxidative responses and quantum use efficiency in Z. noltii were explored through multifactorial analysis.
- Periodicity of propagule expulsion and settlement in the competing native and invasive brown seaweeds, Cystoseira humilis and Sargassum muticum (Phaeophyta)Publication . Engelen, Aschwin; Espirito-Santo, C.; Simões, T.; Monteiro, C. A.; Serrão, Ester; Pearson, G. A.; Santos, RuiDense stands of the invasive species Sargassum muticum (Yendo) Fensholt develop in tidal pools close to its southern distributional limit in Europe, the southwest coast of Portugal. Along this coast, sheltered tidal pools form a specific habitat in which colonization occurs. Invaded pools are originally inhabited by Cystoseira humilis Kützing. Differences in gamete release between the competing native and alien species might be important for the initial settlement and further spread of the invader. Therefore, we tested whether egg expulsion and embryo settlement in both species had the same timing with respect to lunar and tidal cycles. For more than 2 months during the reproductive season egg expulsion and embryo release were monitored daily for each species. Egg expulsion in S. muticum showed a broadly semilunar periodicity peaking around full and new moon (spring tides), when low tides take place in the morning/evening. In contrast, C. humilis egg expulsion showed an asymmetric semilunar-to-lunar periodicity peaking around waning quarter moon, when low tides occur around midday. Embryo settlement detected in pools was low for both species and less periodic. Phase differences in expulsion events between the two species with respect to the semilunar cycle suggest that cues other than the moon are involved in their timing. Our observations suggest that variations in physiological mechanisms and/or environmental conditions result in different patterns of egg expulsion between the two species. This might have consequences for fertilization success, gamete dispersal and survival. It was further found that peaks in egg expulsion and embryo release (i.e. settlement) in S. muticum were much more synchronous at a site in northern Portugal compared with a site close to the current southern distributional limit in south-west Portugal, possibly as a consequence of thermal stress experienced in the south.