Giulia Ceccherelli

Spread of introduced Caulerpa species in macroalgal habitats

A short-term field experiment was designed to identify layers of Mediterranean macroalgal assemblage conducive to successful spread of two introduced Caulerpa species (Bryopsidales, Chlorophyta). By manipulation of species presence, three experimental assemblages were obtained: (1) encrusting algae, having removed the turf and erect species; (2) encrusting and turfing algae, having removed erect species; (3) encrusting, turfing and erect algae, that is, unmanipulated assemblages, which served as a control. Fragments of the two introduced species Caulerpa taxifolia (Vahl) C. Agardh and Caulerpa racemosa (Forsskal) J. Agardh were transplanted in each of the three assemblages. Width of the colony, blade density and percentage of the substratum covered by the two species were measured. The susceptibility of the indigenous community to the spread of Caulerpa species was related to type of assemblage. Blade density and amount of substratum covered by the two Caulerpa species were different between species and generally greater for C. taxifolia than for C. racemosa. Overall, the spread of these species was strongly dependent on the type but not directly on the complexity of the assemblage. Turf was more favourable than encrusting species alone, while the least advantageous habitat was where the macroalgal assemblage is composed of encrusting, turf and erect species. In other words, increased number of species in the assemblage reduces invasion of the Caulerpa species but the type of algae in the assemblage is likely to be more important than number of species. The presence of turf promotes the spread of Caulerpa species.

Response of the non-indigenous Caulerpa racemosa (Forsskål) J. Agardh to the native seagrass Posidonia oceanica (L.) Delile: effect of density of shoots and orientation of edges of meadows

Abstract Caulerpa racemosa is a tropical green alga introduced into the Mediterranean as an immigrant from the Red Sea which has successfully fast-spread in the south-eastern and in the north-western part of the basin. C. racemosa occurs mostly in shallow but also in deep subtidal habitats colonising hard and soft substrata where turfs, erect algae and even seagrasses are present with the potential to profoundly alter indigenous communities. However, the extent to which biotic interactions influence the spread of the alga is not well known. In this study the effects of the presence of the native seagrass Posidonia oceanica on the non-indigenous alga Caulerpa racemosa are examined: a multifactorial experiment was designed to test (1) the importance of the seagrass canopy structure and (2) of orientation of seagrass edge on algal performance along the edge and inside the meadow of P. oceanica and (3) whether patterns of algal growth are consistent at different spatial scales (few centimetres to several metres). The aim of this study is to provide a basis for further experimental investigations of the factors and mechanisms affecting the performance of this alga in the Mediterranean. The results of this study indicated that where Caulerpa racemosa is at the edge of Posidonia oceanica meadow, the vertical growth of the alga (blade length) is sensitive to the combination of time, seagrass density and edge-meadow orientation, that the spread of the alga along the edge of the seagrass meadow is dependent on the characteristics of the area and that the growth of the alga inside the meadow is influenced by seagrass density. The findings of this study suggest that the susceptibility of the indigenous P. oceanica community to invasion of the introduced alga C. racemosa is related to the availability of sand habitat ground created, since low invasion of the very dense edges of the seagrass was observed compared to the less dense ones.

Nutrient availability in the sediment and the reciprocal effects between the native seagrass Cymodocea nodosa and the introduced rhizophytic alga Caulerpa taxifolia

Two reciprocal experiments testing for the effects of nutrient addition in the sediment and competitive interactions between the native seagrass Cymodocea nodosa (Ucria) Ascherson and the introduced alga Caulerpa taxifolia (Vahl) C. Agardh were performed. This study was conducted for 13 months (August 1995 until September 1996) in a bay on the south coast of Elba Island (Italy). Each experiment consisted of the manipulation of the level of nutrients (addition vs. control) and the manipulation of the neighbours (presence vs. removal). Response variables were blade density and size for one experiment and shoot density and leaf length of seagrass in the other. Results indicated that the presence of Caulerpa taxifolia did not affect significantly Cymodocea nodosa shoot density and the increased nutrient availability in the sediment did not alter this pattern. Neither the removal of the canopy of the seagrass nor the fertilization of the sediment has influenced significantly the density of the alga. Both species, where co-occurring, show larger size than where the neighbour is removed. Hence, results of this study suggest that the two species on the long term are likely to coexist and that the high nutrient supply of the sediment would not enhance the probability of success neither of the seagrass nor of the alga. Predictions made on the basis of short-term results, that high nutrient loads of the substratum would have represented an even more suitable condition for C. taxifolia to colonize C. nodosa beds and that on the long-term the alga has a high probability of success, did not occur.

Comparative study of the growth of the two co-occurring introduced green algae Caulerpa taxifolia and Caulerpa racemosa along the Tuscan coast (Italy, western Mediterranean)

Abstract Caulerpa taxifolia and C. racemosa are tropical green algae now found in the Mediterranean, where the former was accidentally introduced and the latter presumably also introduced; both species have spread rapidly and invaded a variety of habitats. Since 1996, the two algae have co-inhabited a region along the Tuscan coast near Livorno. The present study has been carried out to evaluate the invasive capacity of the two species. After one year, the increase in surface area was 67.5 % for C. taxifolia and 284.8 % for C. racemosa , while the increase in patch number was 11.4 % and 121.4 % respectively. The stolon growth of the two algae followed a similar trend, as is showing the same temporal variations with the maximum growth rate between August and October. At the end of the study period, the increase in stolon length and number was significantly higher in C. racemosa than in C. taxifolia . The results of the present study show a higher invasive potential in C. racemosa than in C. taxifolia , although further studies are necessary to evaluate the competition between the two algae and the effect of their simultaneous presence on the benthic community.

Caulerpa racemosa var. cylindracea as a potential source of organic matter for benthic consumers: evidences from a stable isotope analysis

Caulerpa racemosa var. cylindracea has been invading various types of substrates in wide areas throughout the Mediterranean Sea. However, the effects of the distribution of this alga on zoobenthos are scanty. The aim of this study was to evaluate the effects of C. racemosa invasion on the feeding habits of some zoobenthic taxa inhabiting the upper infralittoral rocky shores. This was done by identifying the isotopic N and C ratios of several potential food sources and testing differences in isotope composition among the taxa collected from areas invaded and not-invaded areas by C. racemosa. Results suggest that C. racemosa detritus was a significant food source for the polychaete Syllis prolifera, the gammarid Corophium sextonae and the gastropods Cerithium rupestre and Pisinna glabrata. They would also suggest that stable carbon and nitrogen isotope ratios deserve further attention as a possible valuable approach to understand the overall effect of C. racemosa spread on the trophic interactions among the species.

Habitat and Scale Shape the Demographic Fate of the Keystone Sea Urchin Paracentrotus lividus in Mediterranean Macrophyte Communities

Demographic processes exert different degrees of control as individuals grow, and in species that span several habitats and spatial scales, this can influence our ability to predict their population at a particular life-history stage given the previous life stage. In particular, when keystone species are involved, this relative coupling between demographic stages can have significant implications for the functioning of ecosystems. We examined benthic and pelagic abundances of the sea urchin Paracentrotus lividus in order to: 1) understand the main life-history bottlenecks by observing the degree of coupling between demographic stages; and 2) explore the processes driving these linkages. P. lividus is the dominant invertebrate herbivore in the Mediterranean Sea, and has been repeatedly observed to overgraze shallow beds of the seagrass Posidonia oceanica and rocky macroalgal communities. We used a hierarchical sampling design at different spatial scales (100 s, 10 s and <1 km) and habitats (seagrass and rocky macroalgae) to describe the spatial patterns in the abundance of different demographic stages (larvae, settlers, recruits and adults). Our results indicate that large-scale factors (potentially currents, nutrients, temperature, etc.) determine larval availability and settlement in the pelagic stages of urchin life history. In rocky macroalgal habitats, benthic processes (like predation) acting at large or medium scales drive adult abundances. In contrast, adult numbers in seagrass meadows are most likely influenced by factors like local migration (from adjoining rocky habitats) functioning at much smaller scales. The complexity of spatial and habitat-dependent processes shaping urchin populations demands a multiplicity of approaches when addressing habitat conservation actions, yet such actions are currently mostly aimed at managing predation processes and fish numbers. We argue that a more holistic ecosystem management also needs to incorporate the landscape and habitat-quality level processes (eutrophication, fragmentation, etc.) that together regulate the populations of this keystone herbivore.

Harvesting Effects on Paracentrotus lividus Population Structure: A Case Study from Northwestern Sardinia, Italy, before and after the Fishing Season

Abstract PAIS, A.; SERRA, S.; MELONI, G.; SABA, S., and CECCHERELLI, G., 2012. Harvesting effects on Paracentrotus lividus population structure: A case study from northwestern Sardinia, Italy, before and after the fishing season. The edible sea urchin Paracentrotus lividus is commercially exploited in many European countries. In recent years, however, its stocks have shown a large reduction in several regions because of overfishing. This paper reports the effects of harvesting on P. lividus population structure in shallow rocky reefs from a central-western Mediterranean area where this activity is intensively practiced. Density, size (test diam), and biomass of sea urchin were assessed in October 2004 (just before the fishing season) and in May 2005 (immediately after the fishing season) at two locations where intense P. lividus harvest occurs and at two controls within an adjacent Marine Protected Area (MPA) where P. lividus harvest is prohibited. Density, size, and average biomass of P. lividus were significantly higher at the protected areas than at the exploited ones. In particular, large-sized specimens (test diam >5 cm) were notably less abundant at the fished locations; however, very similar results were detected for both the periods examined (i.e., before and after the fishing season), suggesting that effects of pressing activity of harvest through the past years may have produced effects much more evident than those attributable to the single fishing season. These results reveal the existence of a heavy-fishing impact on P. lividus in northwestern Sardinia and the urgent need of regulating its harvesting to prevent severe direct effects on its populations.

On the Movement of the Sea Urchin Paracentrotus lividus Towards Posidonia oceanica Seagrass Patches

ABSTRACT The movement of the sea urchin Paracentrotus lividus from a rocky habitat to patches of Posidonia oceanica was investigated with respect to the size of the sea urchins and their location from the edge of the patch. With this aim, a manipulative experiment was conducted (4 times) at a location where several P. oceanica patches were interspersed on rocky platforms. Each time, after an accurate removal of the sea urchins populating them, 15 of these patches were randomly assigned in sets of 3 to 5 different urchin addition treatments, using groups of 10 large or small P. lividus specimens (test diameter >50 and <30 mm, respectively) positioned at close and far distances (25 and 100 cm, respectively) from the edges of the patches as follows: large-close, large-far, small-close, small-far, and control patches where no urchins were added. The abundance of sea urchins inside the patches was counted after 24 h. Results highlighted significant variability because of the distance from the patches, whereas no significant effect was observed for sea urchin size. These results suggested that: P. lividus specimens close to P. oceanica patches might have a greater probability of reaching them, and that the chance to reach the patch does not depend on the size of sea urchins. In fact, a comparable ability to move towards the patches was evident for different-sized specimens, indicating that migration from one habitat to the other is possible even for small-sized individuals.

Immediate effects of experimental human trampling on mid-upper intertidal benthic invertebrates at the Asinara Island MPA (NW Mediterranean)

To assess potential risks of human visitation to ecological communities, the immediate effects of human trampling were investigated experimentally on small invertebrates inhabiting mid-upper intertidal hard bottoms covered by algae. Two different experimental intensities of trampling (60 and 120 footsteps) and controls (with no trampling) were applied to quadrats 20 × 20 cm in size (experimental area), within the two ‘no-entry, no-take’ zones of the Asinara Island MPA (Italy, Mediterranean Sea). One day after trampling ended, samples of benthic fauna were collected and the animals attributed to macrofaunal and meiofaunal components. Analyses of variance on the nine most common taxa of macrofauna identified significant higher abundance of bivalves, gammarid amphipods, polychaetes, isopods, oligochaetes in controls than in trampled plots. For nematodes, polychaetes, ostracods, oligochaetes, bivalves, acari, caprellid amphipods and tanaids a significant higher abundance of meiofaunal animals was found in controls than in trampled areas. Although no information on recovery is available, these results suggest that macrofaunal and meiofaunal taxa are vulnerable to this type of disturbance.

Human trampling effects on Mediterranean coastal dune plants

Coastal habitats are particularly vulnerable to recreational impacts because these environments are highly dynamic and continually change in response to biotic and abiotic factors. Sand dune communities are worldwide characterized by high levels of biodiversity, but are often affected by human-induced impacts as those caused by tourist trampling. To understand the effects of human frequentation, trampling, and other human-induced impacts, fencing experiments have been traditionally carried out on coastal dunes. Since in touristic areas dune systems are subjected to different intensities of human frequentations rather than to opening or fencing, in this study we explore the effects of accessibility on vascular plants cover. This study tests the hypothesis that human frequentation on beaches affects spatio-temporal variability of vascular plant abundance on dunes by comparing the plant assemblages of high and low accessible sites in North-East Sardinia (Italy). Our results show that accessibility plays a crucial role in conditioning the percentage of vegetation cover in Mediterranean dunes. In fact, not only we found a perennial vegetation cover that was significantly higher in the sites with low accessibility (and consequently low frequentation), but we also showed that at the sites with high accessibility there were significant differences in vegetation cover between times of sampling (cover was higher before than after summer): on the contrary, differences in perennial vegetation cover among times were not significant at the low frequentation sites. After summer, the difference among low and high frequentation sites in species composition and cover was >90%. Multivariate analysis identified those species that play a pivotal role in differentiating the low and the high frequentation sites. Among them, Crucianella maritima and Sporobolus virginicus can be considered as differential species. Overall, our data show vegetation and plant species responses to human-induced impacts, and are therefore important to support conservation actions in Mediterranean coastal areas interested by mass tourism.