Variation in multiple traits of vegetative and reproductive seagrass tissues influences plant–herbivore interactions

Plant–herbivore interactions have strong ecological and evolutionary consequences, but have been traditionally overlooked in marine higher plants. Despite recent advances in seagrass ecology that highlight the importance of herbivory, the mechanisms that regulate the feeding behaviour of seagrass consumers remain largely unknown. Herbivores have been shown to reduce the sexual reproductive success of seagrasses through direct consumption of inflorescences and seeds, but we know little about intraspecific variation in susceptibility to grazing of different seagrass tissues. We contrasted the relative palatability of reproductive and vegetative tissues of the temperate seagrass Posidonia oceanica in the field, and we assessed the feeding preferences among these tissues of the main consumers of the plant, the fish Sarpa salpa and the urchin Paracentrotus lividus. Moreover, we identified the plant traits that explained the observed feeding behaviour. We provide strong evidence for herbivore selectivity among seagrass tissues. In the field, 70–90% of inflorescences were damaged by herbivores compared to 3–60% of leaves of similar age. In feeding assays, the urchin P. lividus showed over a twofold preference for reproductive tissue at various stages of development. By contrast, we detected no feeding activity on either leaves or inflorescences from the fish S. salpa, which is known to migrate to deeper waters soon after flowering starts and during the period of fruit maturation. Despite being the preferred food of urchins, inflorescences were chemically defended, had higher levels of phenolics and lower nutrient and calorific content than leaves. We experimentally demonstrated that leaf structural defences are the primary factor in determining urchin feeding preferences. Removal of plant structure results in a drastic shift in urchin selectivity towards the most nutritious and less chemically defended leaf tissue, indicating that multiple mechanisms of defence to herbivory may coexist in seagrasses.     

Patterns of fish and sea urchin grazing on tropical Indo-Pacific seagrass beds

Thalassodendron ciliatum shoots were collected from natural populations of fished and unfished protected seagrass meadows to assess the herbivory of fish and sea urchins. Fish herbivory was restricted to unfished seagrass meadows, while sea urchin herbivory took place in fished as well as unfished areas where fish appeared not to be effective urchin predators. The results of this study confirm that grazing by sea urchins is important in tropical seagrass ecosystems and indicate that herbivorous fish graze, and probably consume, substantial amounts of seagrass production in fishing-protected habitats. The fact that much of the information on seagrass herbivory comes from heavily-fished meadows indicates that large-scale studies, which include unfished areas, are necessary to provide reliable spatial patterns of seagrass grazing distribution.     

The effects of sea urchin grazing and drift algal blooms on a subtropical seagrass bed community

Subtropical seagrass beds can be subject to relatively high levels of direct herbivory and large blooms of drift algae, both of which can have important effects on the floral and faunal components of the community. Caging experiments were used to investigate these factors in a Thalassia testudinum bed in Biscayne Bay, Florida. Abundance of sea urchins, Lytechinus variegatus, and drift algae was manipulated within the cages. Naturally occurring levels of urchin grazing do not appear to affect the T. testudinum population. With experimentally increased urchin densities in the winter, seagrass shoot density and aboveground biomass decreased significantly. Similar effects were not detected in the summer, indicating that the impact of grazing on T. testudinum is lessened during this time of year. Shoot density was more vulnerable to grazing than aboveground biomass. This may be a result of grazing-induced increases in seagrass productivity, in which the remaining shoots produce more or longer leaves. In the winter, drift algal blooms form large mats that cover the seagrass canopy. Under the normal grazing regime these algal blooms do not have significant negative effects on the seagrass. With increased grazing pressure, however, there is a synergistic effect of grazing and drift algae on seagrass shoot density. At intermediate urchin density (10 per m(-2)), cages without algae did not undergo significant decreases in shoot density, while those with algae did. At the high density of urchins, the number of seagrass shoots in cages both with and without algae decreased, but the effect was more pronounced for cages with algae. Invertebrate abundance at the field site was low relative to other seagrass beds. There were no discernible effects, either positive or negative, of urchin and algae manipulations on the sampled invertebrate community.     

Disturbance and recovery following catastrophic grazing: studies of a successional chronosequence in a seagrass bed

In August 1997, a large aggregation of the common sea urchin, Lytechinus variegatus , was discovered moving southward through a lush and productive seagrass monoculture of Syringodium filiforme in the Florida Keys, FL. Sea urchin densities at the grazing front were greater than 300 individuals m⁻² which resulted in the overgrazing of seagrasses and a complete denuding of all vegetation from this area. The steady rate of the grazing front migration permitted the estimation of the time since disturbance for any point behind this grazing front allowing the use of a chronosequence approach to investigate the processes early on in succession of these communities.
In May 1999, six north-south parallel transects were established across the disturbed seagrass communities and into the undisturbed areas south of the grazing front. Based on the measured rates of the migration of the grazing front, we grouped 60 sites into five categories (disturbed, recently grazed, active grazing front, stressed and undisturbed). The large scale loss of seagrass biomass initiated community-wide cascading effects that significantly altered resource regimes and species diversity. The loss of the seagrass canopy and subsequent death and decay of the below-ground biomass resulted in a de-stabilization of the sediments. As the sediments were eroded into the water column, turbidity significantly increased, reducing light availability and significantly reducing the sediment nitrogen pool and depleting the seed bank. The portion of the chronosequence that has had the longest period of recovery now consists of a mixed community of seagrass and macroalgae, as remnant survivors and quick colonizers coexist and jointly take advantage of the open space.     

Experimental evidence that herbivory increases shoot density and productivity in a subtropical turtlegrass (Thalassia testudinum ) meadow

The sea urchin, Lytechinus variegatus, has been estimated to consume between 50 and 90% of annual net aboveground production in selected turtlegrass (Thalassia testudinum) meadows in the eastern Gulf of Mexico. Nevertheless, turtlegrass persists where sea urchin grazing is intense. We hypothesized that turtlegrass productivity is stimulated by grazing, as has been reported from terrestrial grassland systems, and that this best explains the persistence of heavily grazed turtlegrass in St. Joseph Bay, Fla. This hypothesis was tested by manipulating sea urchin densities (0, 10, and 20 individuals/m²) in 1-m² enclosures. These densities encompass the range of average densities at the study site and those reported in the literature. Changes in turtlegrass abundance (e.g., short shoot density and biomass), production by short shoots, and leaf width were monitored in these enclosures during the summer. Repeated-measures ANOVA showed that production by short shoots, leaf density/shoot, and leaf width decreased over time in all treatments. Leaf density/shoot and leaf width were not significantly impacted by grazing, nor did grazing significantly reduce seagrass biomass, but it did lead to significantly higher densities of short shoots than found in control cages. Our results indicate that turtlegrass compensates for the effects of sea urchin herbivory by increasing the recruitment of short shoots during the growing season. We estimate that this increased shoot density led to a 40% increase in net aboveground primary production (g dry weight/m²) in grazing treatments, which helps to explain the lack of significant reductions of sea grass biomass during the growing season. Received: 26 April 1995 / Accepted: 29 May 1997     

The role of the sea urchin,Tripneustes gratilla (Linnaeus), in decomposition and nutrient cycling in a tropical seagrass bed

The sea urchin,Tripneustes gratilla, which feeds mainly on living leaves of the seagrass,Thalassia hemprichii, was studied in its habitat on the southern coast of Papua New Guinea, and its roles in decomposition and nutrient cycling in a seagrass bed were assessed through the excretion of ammonium and metabolism of feces produced by the sea urchin. Carbon content of the fresh feces (21% of dry weight) was similar to that of intact dead leaves of the same species (22–23%). Carbon/nitrogen and carbon/phosphorus ratios of the feces (21.7 and 466, respectively), however, were significantly lower than those of the dead leaves (25.9–27.7 and 656–804, respectively), indicating that the feces retain more nitrogen and phosphorus in comparison with carbon. Net consumption of ammonium and orthophosphate typically concurred with oxygen consumption during dark incubation of both the dead leaves and the sea urchin feces. Compared with the same oxygen consumption rate, however, the dead leaves consumed more orthophosphate than the feces. Under sunlight, dead leaves showed a net accumulation of carbon by epiphytic algae, while the feces showed a carbon loss. Ammonium excretion by this sea urchin (1.7–5.4 mg nitrogen/individual/day) would thus appear to make a significant contribution to nitrogen recycling since biological communities associated with dead leaves and sea urchin feces tend to demand an external supply of nitrogen, such as ammonium.     

Herbivore control in connected seascapes: habitat determines when population regulation occurs in the life history of a key herbivore

Herbivore outbreaks often trigger catastrophic overgrazing events in marine macrophyte ecosystems. The sea urchin Paracentrotus lividus, the dominant herbivore of shallow Mediterranean seascapes, is capable of precipitating shifts to barrens when its populations explode. Paracentrotus lividus is found ubiquitously in rocky macroalgal communities and in sandy seagrass meadows of Posidonia oceanica, two of the most important subtidal habitats in the Mediterranean. We explored if habitat‐specific regulation across the principal stages of the urchin life cycle could help explain the persistence of these populations in connected mosaics. We measured each of three relevant ecological processes (i.e. settlement, post‐settlement survival and predation) across a wide stretch of the Mediterranean coast (ca 600 km). Our results show that habitat‐specific regulation is critical in determining urchin populations: each habitat limited urchin sub‐populations at different life stages. Settlement was never limiting; urchins settled at similar rates in both habitats across the coast. Post‐settlement survival was a clear bottleneck, particularly in seagrass meadows where no juvenile urchins were recorded. Despite this bottleneck in seagrasses, adult urchin populations were very similar in both seagrass and macroalgal habitats indicating that other processes (potentially migration) could be key in determining adult distributions across the mosaic. The fact that population regulation is clearly habitat‐specific suggests that sea urchin populations may be significantly buffered from bottlenecks in mixed seascapes where both habitats co‐occur. Sea urchin populations can therefore persist across the seascape despite strong habitat‐specific regulation either by maintaining reproductive output in one habitat or by migrating between them. By affording these regulatory escapes to habitat‐modifying species, patchy mosaics may be much more prone to herbivore outbreaks and a host of cascading effects that come in their wake.     

Accounting for size‐specific predation improves our ability to predict the strength of a trophic cascade

Predation can influence the magnitude of herbivory that grazers exert on primary producers by altering both grazer abundance and their per capita consumption rates via changes in behavior, density‐dependent effects, and size. Therefore, models based solely on changes in abundance may miss key components of grazing pressure. We estimated shifts in grazing pressure associated with changes in the abundance and per capita consumption rates of sea urchins triggered by size‐selective predation by sea otters (Enhydra lutris). Field surveys suggest that sea otters dramatically decreased the abundance and median size of sea urchins. Furthermore, laboratory experiments revealed that kelp consumption by sea urchins varied nonlinearly as a function of urchin size such that consumption rates increased to the 0.56 and 0.68 power of biomass for red and green urchins, respectively. This reveals that shifts in urchin size structure due to size‐selective predation by sea otters alter sea urchin per capita grazing rates. Comparison of two quantitative models estimating total consumptive capacity revealed that a model incorporating shifts in urchin abundance while neglecting urchin size structure overestimated grazing pressure compared to a model that incorporated size. Consequently, incorporating shifts in urchin size better predicted field estimates of kelp abundance compared to equivalent models based on urchin abundance alone. We provide strong evidence that incorporating size‐specific parameters increases our ability to describe and predict trophic interactions.     

Trophic Cascades Induced by Lobster Fishing Are Not Ubiquitous in Southern California Kelp Forests

Fishing can trigger trophic cascades that alter community structure and dynamics and thus modify ecosystem attributes. We combined ecological data of sea urchin and macroalgal abundance with fishery data of spiny lobster (Panulirus interruptus) landings to evaluate whether: (1) patterns in the abundance and biomass among lobster (predator), sea urchins (grazer), and macroalgae (primary producer) in giant kelp forest communities indicated the presence of top-down control on urchins and macroalgae, and (2) lobster fishing triggers a trophic cascade leading to increased sea urchin densities and decreased macroalgal biomass. Eight years of data from eight rocky subtidal reefs known to support giant kelp forests near Santa Barbara, CA, USA, were analyzed in three-tiered least-squares regression models to evaluate the relationships between: (1) lobster abundance and sea urchin density, and (2) sea urchin density and macroalgal biomass. The models included reef physical structure and water depth. Results revealed a trend towards decreasing urchin density with increasing lobster abundance but little evidence that urchins control the biomass of macroalgae. Urchin density was highly correlated with habitat structure, although not water depth. To evaluate whether fishing triggered a trophic cascade we pooled data across all treatments to examine the extent to which sea urchin density and macroalgal biomass were related to the intensity of lobster fishing (as indicated by the density of traps pulled). We found that, with one exception, sea urchins remained more abundant at heavily fished sites, supporting the idea that fishing for lobsters releases top-down control on urchin grazers. Macroalgal biomass, however, was positively correlated with lobster fishing intensity, which contradicts the trophic cascade model. Collectively, our results suggest that factors other than urchin grazing play a major role in controlling macroalgal biomass in southern California kelp forests, and that lobster fishing does not always catalyze a top-down trophic cascade.     

Plant-herbivore interactions in seagrass meadows

During the past two decades we have gained much insight into the factors that regulate the productivity of seagrass dominated ecosystems, especially those at low latitudes. Here, we review and reassess the importance of plant-herbivore interactions in seagrass meadows, focusing on recent studies that have examined: 1) grazing on live seagrass leaves; 2) consumption of epiphytic algae growing on seagrass leaves; and 3) consumption of planktonic algae from the waters surrounding seagrass meadows. The major conclusion is that, in contrast to what has been reported in much of the literature on food webs in seagrass meadows, a diverse grazing pathway continues to represent an important conduit for the transfer of energy from the primary producers to higher order consumers. This remains true, although in many areas consumption of seagrasses is reduced in an historical context, owing to the overharvesting of many large species of herbivorous waterfowl, turtles and mammals.We also summarize our view of the important gaps in understanding the broadly defined topic of herbivory in seagrass-dominated ecosystems. We suggest that future studies should focus on: understanding the foraging strategies of seagrass herbivores; quantifying the impact of herbivory on seagrass demography, including effects on sexual reproduction, the fate of flowers, and the production of fruits and seeds; and documenting the commonness of compensatory responses to grazing. In addition, the role of chemical defenses in seagrass species remains inadequately investigated. Studies of the roles of nutritional content (as measured by C/N/P ratios) and chemical defenses are also fertile grounds for future studies of epiphytes and their grazers, as are additional experiments to quantify the relative roles of top-down and bottom-up factors as they determine algal growth and abundance. There is also a need to expand the geographical scope of studies of epiphyte-grazer interactions from cold temperate to sub-tropical and tropical waters. Suspension feeders also need to be studied more broadly, with additional experiments required to quantify their effects on water clarity and their ability to fertilize pore waters, and whether benefits from these activities balances the costs of shading and competition for space that can result from both epifaunal and infaunal suspension feeders.     

Specificity in Mesograzer-Induced Defences in Seagrasses

Grazing-induced plant defences that reduce palatability to herbivores are widespread in terrestrial plants and seaweeds, but they have not yet been reported in seagrasses. We investigated the ability of two seagrass species to induce defences in response to direct grazing by three associated mesograzers. Specifically, we conducted feeding-assayed induction experiments to examine how mesograzer-specific grazing impact affects seagrass induction of defences within the context of the optimal defence theory. We found that the amphipod Gammarus insensibilis and the isopod Idotea chelipes exerted a low-intensity grazing on older blades of the seagrass Cymodocea nodosa, which reflects a weak grazing impact that may explain the lack of inducible defences. The isopod Synischia hectica exerted the strongest grazing impact on C. nodosa via high-intensity feeding on young blades with a higher fitness value. This isopod grazing induced defences in C. nodosa as indicated by a consistently lower consumption of blades previously grazed for 5, 12 and 16 days. The lower consumption was maintained when offered tissues with no plant structure (agar-reconstituted food), but showing a reduced size of the previous grazing effect. This indicates that structural traits act in combination with chemical traits to reduce seagrass palatability to the isopod. Increase in total phenolics but not in C:N ratio and total nitrogen of grazed C. nodosa suggests chemical defences rather than a modified nutritional quality as primarily induced chemical traits. We detected no induction of defences in Zostera noltei, which showed the ability to replace moderate losses of young biomass to mesograzers via compensatory growth. Our study provides the first experimental evidence of induction of defences against meso-herbivory that reduce further consumption in seagrasses. It also emphasizes the relevance of grazer identity in determining the level of grazing impact triggering resistance and compensatory responses of different seagrass species.     

Population of Lytechinus variegatus (Echinoidea: Toxopneustidae) and structural characteristics of seagrass of Thalassia testudinum in Mochima Bay,Venezuela)

To compare the general features of Thalassia testudinum seagrass at Mochima Bay with sea urchin (Lxtechinus variegatus) abundance and distribution, three T. testudinum seagrass beds were selected, from the mouth (strong wave exposure) to the inner bay (calm waters). Each site was surveyed by using 5 line transects (20 m long) parallel to the coast and 1 m2 quadrats. In situ measurements of T. testudinum cover, shoot and leaf density were taken. Estimation of dry biomass for each seagrass fraction (leaves, rhizomes and roots) and leaf length were obtained from 25 vegetation samples extracted per site using cores (15 cm diameter). A multivariate analysis of variance (Manova) and a less significative difference test (LSD) were performed to examine differences between sites and within sites at different depths. A stepwise multiple regression analysis was done, dependent variable was sea urchin density; independent variables: vegetation values at each site. The only seagrass species found in the three sites was T. testudinum, and cover was 56-100%, leaf density 100-1000 leaf/m2, lengths 6-18.8 cm and shoot density 20-475 shoots/m2. The highest sea urchin densities were found at Isla Redonda and Ensenada Toporo (1-3.6 ind/m2), the lowest at Playa Colorada (0.6-0.8 ind/m2). Significant differences in seagrass features between sites were obtained (Manova p < 0.001), but not between depths (Manova p < 0.320). The regression coefficient between sea urchin density and seagrass parameters was statistically significant (r2 = 0.154, p < 0.007), however, total biomass was the only variable with a significant effect on sea urchin distribution (beta = 0.308, p < 0.032). The other variables did not explain satisfactorily L. variegatus abundance and distribution.     

Responses of seagrass to anthropogenic and natural disturbances do not equally translate to its consumers

Coastal communities are under threat from many and often co‐occurring local (e.g., pollution, eutrophication) and global stressors (e.g., climate change), yet understanding the interactive and cumulative impacts of multiple stressors in ecosystem function is far from being accomplished. Ecological redundancy may be key for ecosystem resilience, but there are still many gaps in our understanding of interspecific differences within a functional group, particularly regarding response diversity, that is, whether members of a functional group respond equally or differently to anthropogenic stressors. Herbivores are critical in determining plant community structure and the transfer of energy up the food web. Human disturbances may alter the ecological role of herbivory by modifying the defense strategies of plants and thus the feeding patterns and performance of herbivores. We conducted a suite of experiments to examine the independent and interactive effects of anthropogenic (nutrient and CO₂ additions) and natural (simulated herbivory) disturbances on a seagrass and its interaction with two common generalist consumers to understand how multiple disturbances can impact both a foundation species and a key ecological function (herbivory) and to assess the potential existence of response diversity to anthropogenic and natural changes in these systems. While all three disturbances modified seagrass defense traits, there were contrasting responses of herbivores to such plant changes. Both CO₂ and nutrient additions influenced herbivore feeding behavior, yet while sea urchins preferred nutrient‐enriched seagrass tissue (regardless of other experimental treatments), isopods were deterred by these same plant tissues. In contrast, carbon enrichment deterred sea urchins and attracted isopods, while simulated herbivory only influenced isopod feeding choice. These contrasting responses of herbivores to disturbance‐induced changes in seagrass help to better understand the ecological functioning of seagrass ecosystems in the face of human disturbances and may have important implications regarding the resilience and conservation of these threatened ecosystems.     

Habitat-dependent growth in a Caribbean sea urchin <Emphasis Type="Italic">Tripneustes ventricosus</Emphasis>: the importance of food type

The sea urchin Tripneustes ventricosus is a common, yet relatively poorly known, grazer of seagrass beds and coral reefs throughout the Caribbean. We compared the size and abundance of urchins between adjacent seagrass and coral reef habitats (where macroalgae are the dominant primary producers). We also conducted a laboratory experiment comparing the growth rate of juvenile urchins fed a diet of either macroalgae or seagrass. Reef urchins had significantly larger test diameter than those in the seagrass on some sampling dates. This size difference may be at least partially explained by diet, because laboratory-reared urchins fed macroalgae grew significantly faster than those fed seagrass. The seagrass population, however, was stable over time, whereas the reef population exhibited strong fluctuations in abundance. Overall, our study indicates that both the seagrass and coral reef habitats are capable of supporting healthy, reproductive populations of T. ventricosus. Each, however, appears to offer a distinct advantage: faster growth on the reef and greater population stability in the seagrass.     

Effects of sediment nutrients on seagrasses: Literature review and mesocosm experiment

A review of the literature shows that seagrass growth, abundance and morphology are strongly linked to available nutrient resources. In north temperate climates and in habitats with terrigenous sediments, nitrogen limitation occurs in both intertidal and subtidal beds. Typically, seagrasses growing in terrigenous sediments have ample phosphorus but lack nitrogen, and the plants' chemical composition is depleted in nitrogen. However, seagrasses occurring in tropical environments and carbonate sediments appear to experience phosphorus limitation due to binding of phosphate in the sediments. Thus, it is the sediment geochemistry in seagrass beds that is important in determining the limiting nutrient to seagrass growth.Examination of the literature indicates that field research on seagrass relationships involves too many interactive factors to be able to say certainly that any one plant characteristics is caused by any one environmental factor. Using mesocosms (partially enclosed outdoor experimental set-ups) one environmental factor can be changed between the treatment tank and the control. Therefore, we can determine experimentally that a plant characteristic is affected by a particular altered environmental factor. Experimental mesocosms used to grow eelgrass, Zostera marina L., in substrata of varied nitrogen composition showed the dramatic effect of insufficient nitrogen on eelgrass growth, abundance and leaf morphology. Additionally, eelgrass leaf tissue from low ammonium sand substratum is significantly depleted in nitrogen, demonstrating the supposition that nitrogen is limiting in terrigenous environments.     

Echinoderms display morphological and behavioural phenotypic plasticity in response to their trophic environment

The trophic interactions of sea urchins are known to be the agents of phase shifts in benthic marine habitats such as tropical and temperate reefs. In temperate reefs, the grazing activity of sea urchins has been responsible for the destruction of kelp forests and the formation of 'urchin barrens', a rocky habitat dominated by crustose algae and encrusting invertebrates. Once formed, these urchin barrens can persist for decades. Trophic plasticity in the sea urchin may contribute to the stability and resilience of this alternate stable state by increasing diet breadth in sea urchins. This plasticity promotes ecological connectivity and weakens species interactions and so increases ecosystem stability. We test the hypothesis that sea urchins exhibit trophic plasticity using an approach that controls for other typically confounding environmental and genetic factors. To do this, we exposed a genetically homogenous population of sea urchins to two very different trophic environments over a period of two years. The sea urchins exhibited a wide degree of phenotypic trophic plasticity when exposed to contrasting trophic environments. The two populations developed differences in their gross morphology and the test microstructure. In addition, when challenged with unfamiliar prey, the response of each group was different. We show that sea urchins exhibit significant morphological and behavioural phenotypic plasticity independent of their environment or their nutritional status.     

Effects of invasive seaweeds on feeding preference and performance of a keystone Mediterranean herbivore

The consequences of invasive species on ecosystem processes and ecological interactions remain poorly understood. Predator–prey interactions are fundamental in shaping species evolution and community structure and can be strongly modified by species introductions. To fully understand the ecological effects of invasive species on trophic linkages it is important to characterize novel interactions between native predators and exotic prey and to identify the impacts of invasive species on the performance of native predators. Although seaweed invasions are a growing global concern, our understanding of invasive algae—herbivore interactions is still very limited. We used a series of feeding experiments between a native herbivore and four invasive algae in the Mediterranean Sea to examine the potential of native sea urchins to consume invasive seaweeds and the impacts of invasive seaweed on herbivore performance. We found that three of the four invasive species examined are avoided by native herbivores, and that feeding behaviour in sea urchins is not driven by plant nutritional quality. On the other hand, Caulerpa racemosa is readily consumed by sea urchins, but may escape enemy control by reducing their performance. Recognizing the negative impacts of C. racemosa on herbivore performance has highlighted an enemy escape mechanism that contributes to explaining how this widespread invasive alga, which is preferred and consumed by herbivores, is not eradicated by grazing in the field. Furthermore, given the ecological and economic importance of sea urchins, negative impacts of invasive seaweeds on their performance could have dramatic effects on ecosystem function and services, and should be accounted for in sea urchin population management strategies.     

Do native herbivores provide resistance to Mediterranean marine bioinvasions? A seaweed example

Generalist herbivores in marine ecosystems are poorly examined for their potential to serve as a source of biotic resistance against algal invasion. We assessed how one of the main generalist herbivores in Mediterranean rocky reefs (the sea urchin Paracentrotus lividus) affects Lophocladia lallemandii and Caulerpa racemosa, two algal invaders with strong detrimental effects on native benthic communities. In a comparison of sea urchin gut contents to algal community composition, strong preferences were exhibited, leading to no relationship between consumption and availability. Both C. racemosa and L. lallemandi were abundant in algal assemblages (>60% occurrence), but C. racemosa (20% of diet) was consumed more than L. lallemandi (3.5%). Experimental enclosures of sea urchins (12 sea urchins * m⁻²) were carried out in locations where L. lallemandii was already established and C. racemosa was rare (new invasion) or abundant (established invasion). C. racemosa was negatively affected by sea urchins only when it was rare, and no effect was detected when the alga was already abundant. Results for L. lallemandi were exactly opposite: urchins limited seasonal increases in L. lallemandi in highly-invaded areas. Because of the small amount of direct consumption of L. lallemandi, its decrease in abundance may be related to the grazing of native algae where L. lallemandii is attached. Overall, our results show that high densities of native herbivores may reduce invasive algae at low densities, due to a combination of direct and indirect effects, but it has no significant effect in highly-invaded areas.     

Measuring fertilization success of broadcast-spawning marine invertebrates within seagrass meadows

Increasing current velocity has been negatively correlated with the fertilization success of marine broadcast-spawning invertebrates. Seagrass has been shown to affect seawater hydrodynamics by slowing the movement of water. In this study we aimed to tease apart the relationship between fertilization success in sea urchins inside and outside of seagrass beds in St. Joseph Bay, Florida. Fluorescein dye diffusion, as a proxy for gamete diffusion, indicated higher rates of diffusion in sand habitats outside of seagrass beds. We quantified the proportion of eggs that remained on a female compared to being advected off a female over a 2-min interval in and out of grass beds. More eggs were collected inside of seagrass beds than over sand habitats, suggesting increased residence time of gametes within the beds. We induced sea urchins to spawn in experimental arrays in and out of grass beds and measured the fertilization success of eggs released from females and captured in the water column with a plankton pump. The fertilization success of eggs was significantly higher in grass beds. We concluded that seagrasses have the potential to mitigate gamete diffusion and increase the reproductive success of broadcast-spawning species that spawn in them.