Renato Chemello

Trophic cascades in benthic marine ecosystems : lessons for fisheries and protected-area management

An important principle of environmental science is that changes in single components of systems are likely to have consequences elsewhere in the same systems. In the sea, food web data are one of the few foundations for predicting such indirect effects, whether of fishery exploitation or following recovery in marine protected areas (MPAs). We review the available literature on one type of indirect interaction in benthic marine ecosystems, namely trophic cascades, which involve three or more trophic levels connected by predation. Because many indirect effects have been revealed through fishery exploitation, in some cases we include humans as trophic levels. Our purpose is to establish how widespread cascades might be, and infer how likely they are to affect the properties of communities following the implementation of MPAs or intensive resource exploitation. We review 39 documented cascades (eight of which include humans as a trophic level) from 21 locations around the world; all but two of the cascades are from shallow systems underlain by hard substrata (kelp forests, rocky subtidal, coral reefs and rocky intertidal). We argue that these systems are well represented because they are accessible and also amenable to the type of work that is necessary. Nineteen examples come from the central-eastern and north-eastern Pacific, while no well-substantiated benthic cascades have been reported from the NE, CE or SW Atlantic, the Southern Oceans, E Indian Ocean or NW Pacific. The absence of examples from those zones is probably due to lack of study. Sea urchins are very prominent in the subtidal examples, and gastropods, especially limpets, in the intertidal examples; we suggest that this may reflect their predation by fewer specialist predators than is the case with fishes, but also their conspicuousness to investigators. The variation in ecological resolution amongst studies, and in intensity of study amongst systems and regions, indicates that more cascades will likely be identified in due course. Broadening the concept of cascades to include pathogenic interactions would immediately increase the number of examples. The existing evidence is that cascade effects are to be expected when hard-substratum systems are subject to artisanal resource exploitation, but that the particular problems of macroalgal overgrowth on Caribbean reefs and the expansion of coralline barrens in the Mediterranean rocky-sublittoral will not be readily reversed in MPAs, probably because factors other than predation-based cascades have contributed to them in the first place. More cascade effects are likely to be found in the soft-substratum systems that are crucial to so many large-scale fisheries, when opportunities such as those of MPAs and fishing gradients become available for study of such systems, and the search is widened to less conspicuous focal organisms such as polychaetes and crustaceans.

Evaluating the ecological effects of Mediterranean marine protected areas: habitat, scale and the natural variability of ecosystems

Summary The capability to detect and predict the responses of marine populations and communities to the establishment of marine protected areas (MPAs) depends on the ability to distinguish between the influences of management and natural variability due to the effects of factors other than protection. Thus, it is important to understand and quantify the magnitude and range of this natural variability at each scale of observation. Here we review the scale of responses of target populations and communities to protection within Mediterranean MPAs, against their ‘normal’ spatio-temporal heterogeneity, and compare those with documented cases from other temperate and tropical marine ecosystems. Additionally, we approach the problem of the relative importance of habitat structure, considered as a set of biological and physical elements of the seascape hierarchically arranged in space at multiple scales, to drive natural variability. We conclude that much more effort has to be made to characterize heterogeneity in relation to Mediterranean MPAs, and to quantify and explain relationships between target species and their habitats as sources of such variability. These studies should be based on sound sampling designs, which (1) generate long-term data sets, and would ideally (2) be based on a Mediterranean-wide comparison of a number of protected and unprotected localities, (3) be designed from a multi-scaled perspective, and (4) control for factors other than protection, in order to avoid their confounding effects. The need for appropriate spatial and temporal replication, nested designs and power analysis is advocated.

Dendropoma lower intertidal reef formations and their palaeoclimatological significance, NW Sicily

Abstract Most carbonate rocky shores of NW Sicily are marked by a coalescence of shells of the gastropod Dendropoma in a construction that is variably developed as a response to wave impact. Here, we review all the available information on these constructions and find that the fossil reefs are reliable sea-level indicators. The thickness of the reef samples never exceeds 30–40 cm below sea-level, whereas all 14 C dates fall within a range of few centuries. Some small fragments ejected by violent sea storms date back to 2500 years cal BP. No samples older than 6200 years cal BP have been detected so far. The present distribution of Mediterranean vermetid platforms should result from a northward migration related to the long term effect of the Holocene sea surface temperature (SST) warming. Some consideration on the morphology of the reefs and the comparison with the available data point out that Dendropoma reefs are excellent biological indicators of sea-level fluctuations especially when detected and sampled in tectonically stable areas as those in NW Sicily.

Patterns of algal recovery and small-scale effects of canopy removal as a result of human trampling on a Mediterranean rocky shallow community

The ecological importance of marine algae is widely known but in shallow coastal areas the composition and structure of algal communities may be affected by different human activities. Recovery from different trampling disturbances of two competing morphological groups (i.e. macroalgae and algal turfs) and effects of macroalgal canopy removal on the dominant associated fauna were examined using controlled trampling experiments. Six months after trampling disturbance was removed, the two morphological groups closely resembled control (untrampled) conditions, both in terms of cover and canopy (%). In particular, macroalgal recovery seemed to be very rapid: the higher the impact on the system the more rapid the recovery rate. In the short-term, the removal of macroalgal fronds (i.e. canopy) caused evident changes in invertebrate and crypto-benthic fish densities although these indirect effects were species-specific. Erect macroalgae are very sensitive to disturbance and even relatively low intensities of human use may be non-sustainable for this shallow assemblage. The present findings suggest some interesting options for the management of Mediterranean rocky shallow areas. This is crucial for coastal areas that are intended to be maintained in natural condition for conservation purposes.

Ocean acidification impairs vermetid reef recruitment

Vermetids form reefs in sub-tropical and warm-temperate waters that protect coasts from erosion, regulate sediment transport and accumulation, serve as carbon sinks and provide habitat for other species. The gastropods that form these reefs brood encapsulated larvae; they are threatened by rapid environmental changes since their ability to disperse is very limited. We used transplant experiments along a natural CO2 gradient to assess ocean acidification effects on the reef-building gastropod Dendropoma petraeum. We found that although D. petraeum were able to reproduce and brood at elevated levels of CO2, recruitment success was adversely affected. Long-term exposure to acidified conditions predicted for the year 2100 and beyond caused shell dissolution and a significant increase in shell Mg content. Unless CO2 emissions are reduced and conservation measures taken, our results suggest these reefs are in danger of extinction within this century, with significant ecological and socioeconomic ramifications for coastal systems.

Short-term effect of human trampling on the upper infralittoral macroalgae of Ustica Island MPA (western Mediterranean, Italy)

The short-term response of Mediterranean upper infralittoral macroalgal species to experimental human trampling was investigated. Disturbances of six different intensities were applied within the integral reserve of the Ustica Island marine protected area (Italy, Mediterranean Sea). The dominant macroalgal species Cystoseira brachicarpa v. balearica and Dictyota mediterranea were strongly affected by human trampling. Higher levels of disturbance significantly affected both algal percentage cover and canopy at an increasing rate. Three months after trampling, for both variables it was highlighted that the algal recovery from disturbance was incomplete, being significantly different among trampling intensities. The current study revealed that in the short-term it was not possible to identify critical levels of trampling that are sustainable for this shallow community.

Scuba diver behaviour and its effects on the biota of a Mediterranean marine protected area.

The effects of diving activity in different Mediterranean subtidal habitats are scarcely known. This study evaluates diver behaviour (for example time spent in each habitat), use (contacts made with the substrate) and immediate effects of diver contact on benthic species in a marine protected area (MPA) in Sicily. Over a two-year period, intentions of 105 divers were observed within seven subtidal habitats: algae on horizontal substrate, algae on vertical substrate, Posidonia oceanica , encrusted walls, caves, sand and pebbles. Divers selected a habitat in proportion to its availability along the scuba trail. On average, each diver made 2.52 contacts every seven minutes, and no differences were detected among the levels of diver scuba certification. The highest rates of total and unintentional contacts were recorded on caves and encrusted walls, and the slow growing species Eunicella singularis and Astroides calycularis were the most frequently injured by divers. Most of the contacts were concentrated in the first minutes of the dives. The identification of diving effects in different habitats will enable management strategies to specifically control this impact at a habitat scale, for example restricting the start of the dive to low vulnerability habitats would reduce damage to benthic organisms, allowing sustainable use of MPAs.

A new marker for sea surface temperature trend during the last centuries in temperate areas: Vermetid reef

The presence of Vermetid reefs in temperate waters, their diffusion in the Mediterranean Sea, and the possibility of performing 14 C ages allowed the use of Vermetids as an indicator of sea level changes. We present new data on sea climate trend fluctuations that could be interpreted as Sea Surface Temperature (SST) variations, recorded on Vermetid (Dendropoma petraeum) reefs, by means of isotopic analysis. The isotopic records show positive values of the d 18 O relative to present-day values in the period between 1600 and 1850 AD; this deviation occurs in association with the climatic cooling event known as Little Ice Age (LIA). Subsequently, we can observe the warming trend that characterized the last century. These preliminary results indicate that Vermetids could be considered a new SST proxy-data for the Mediterranean Sea and, more generally, for temperate areas. D 2003 Elsevier B.V. All rights reserved.

Molluscan assemblages associated with photophilic algae in the Marine Reserve of Ustica Island (Lower Tyrrhenian Sea, Italy)

Abstract Very few studies have addressed the effect of protection on macrozoobenthos in marine protected areas, and particularly for sites in the Mediterranean Sea. In the present study, the molluscan assemblages associated with photophilic algal communities of the Marine Reserve of Ustica Island, were investigated. A survey was carried out along transects from 1 to 15 m in depth, during the spring of 1996 at three different sites, subjected to different levels of reserve protection. Species richness and number of individuals reflected the level of protection, and both variables were significantly higher in the integral (most heavily protected) part of the reserve compared with the less well protected buffer areas. Species diversity values showed no correlation with the level of protection. The results reported here do not agree with previous findings on the ‘reserve effect’ involving large‐sized macrozoo‐benthic species. However, the data reported, together with a study on polychaetes from the same study sites, and visual census observations concerning the fish assemblage of the island may support the hypothesis that protection of piscivore and macrocar‐nivore species within the integral reserve suppresses populations of small‐sized microcarnivorous fish species, allowing ‘prey‐release’ of small benthic invertebrates.

Vermetid reefs in the Mediterranean Sea as archives of sea-level and surface temperature changes

Vermetid reefs are among the most important bioconstructions in the Mediterranean Sea, with a distribution restricted to the warmest part of the basin. Their structure, and vertical and geographical distribution make them good biological indicators of changes in sea level and sea-surface temperature over the last two millennia.