The spread of Caulerpa racemosa var. cylindracea in the Mediterranean Sea: an example of how biological invasions can influence beta diversity

The present study aims to evaluate if invasion of Caulerpa racemosa var. cylindracea can affect both alpha and beta diversity in Mediterranean coastal system. The following hypotheses were tested: (i) measures of alpha diversity differ between invaded and non-invaded assemblages (ii) this pattern is consistent between different habitats, (iii) assemblages from habitats which are naturally very different tend to be more similar when invaded compared to non-invaded situations. Results showed that Mediterranean benthic assemblages invaded by C. racemosa var. cylindracea were characterized by lower values of alpha diversity than non-invaded assemblages; moreover, differences between deep (25 m) and shallow water (5m) assemblages appeared lower in invaded than in non-invaded areas, with a decrease of beta diversity. This homogenisation of habitats was related to a lack of several characteristic species and to a higher abundance of few opportunistic species in invaded areas, particularly turf-forming algae. Moreover, variability in taxa composition was lower in invaded than in non-invaded assemblages in both habitats.     

Changes in amphipod (Crustacea) assemblages associated with shallow-water algal habitats invaded by Caulerpa racemosa var. cylindracea in the western Mediterranean Sea

The effects of the invasive species Caulerpa racemosa var. cylindracea (hereafter C. racemosa) on amphipod assemblages associated with shallow-water rocky habitats were studied. Two habitats located along the SE Iberian Peninsula were compared; invaded and non-invaded. The results showed that growth of C. racemosa affects habitat structure, influencing the species composition and biomass of macroalgae, and detritus accumulation. In turn, such changes in habitat features affected the associated amphipod assemblages with different ecological requirements. However, the species richness of amphipods was relatively high in both habitats, while the species composition of amphipods changed completely. For example, some species such as Ampithoe ramondi and Hyale schmidti did not colonize invaded habitats, while others such as Apocorophium acutum were favoured by the spread of C. racemosa. Habitat invasion by C. racemosa can have an important influence on biotic assemblages, modifying both habitat structure and the associated fauna, with unknown effects on the overall ecosystem.     

Fragmentation as a Strategy for Caulerpa Species: Fates of Fragments and Implications for Management of an Invasive Weed

Abstract. The successful growth of fragments to full size macroalgal clones is well known for many taxa in mariculture. From an ecological context, however, few studies relate success by invasive species to the ability to fragment readily, disperse widely or re-attach rapidly - obvious hallmarks of weedy species. This study compares morphologically and ecologically distinct species of the green alga Caulerpa : in the Mediterranean, Caulerpa taxifolia is well known as an invasive weed; Caulerpa prolifera grows around or inside seagrass beds and is not apparently invasive; Caulerpa verticillata is among the most diminutive species, with potentially the most restricted reef distribution. We tested three hypotheses: (i) fragment success is independent of size, (ii) fragment success is independent of position within the plant and (iii) fragments from all species will be equally successful in establishing rhizoids, the first step to re-attachment. The success of small fragments (5–40 mm lengths) cut from distinct locations within replicate plants was scored for survival and establishment of rhizoids in bioassays. Additionally, for C. taxifolia , estimates were made of the force needed to puncture upright and prostrate axes, thereby estimating forces required to generate fragments from these two sources. Among the three species, only upright axes of C. taxifolia readily recovered from fragmentation stress for fragment sizes > 10 mm. The forces required to fragment upright axes were substantially less than for comparable rhizome portions. Fragments of C. prolifera were viable only for rhizome portions; fragments of C. verticillata were not viable at any size examined in this study. Estimates of viable fragment size, origin and force required to fragment axes for C. taxifolia support existing field data suggesting that fragmentation is the effective propagation mechanism underlying the spread of this invasive weed.     

Persistence of biological invasion effects: Recovery of macroalgal assemblages after removal of Caulerpa racemosa var. cylindracea

The present study evaluated the capability of recovery of Mediterranean macroalgal assemblages on rocky bottom invaded by Caulerpa racemosa. A manipulative experiment was performed to compare the structure of native assemblages that: (a) were invaded by the alga, (b) were invaded, but where the invasive alga was removed, and (c) had never been invaded. Macroalgal assemblages differed significantly between areas invaded by C. racemosa and control areas. Moreover, one year after removal of the invader, the recovery of community was minimal. Overall, recovery consisted of a significant increase in species richness and cover of macroalgae, but cover of each vegetation layer (encrusting, turf and erect) did not vary significantly between cleared and Invaded plots through time. Results showed that the effects of the colonization persist after the removal of the alga and the process of recovery of the assemblages appears to be quite slow. This finding might be an important contribution to evaluate invasion costs and thus potentially useful for an effective addressing of control strategies.     

Macroalgal assemblages of disturbed coastal detritic bottoms subject to invasive species

Characteristic flora and fauna that are highly sensitive to disturbances colonize coastal detritic bottoms in the Mediterranean Sea. In the present study, a comparison of the assemblage composition and colonization by invasive macroalgae was made between two coastal detritic macrophyte assemblages, one dominated by rhodoliths (free-living non-geniculate Corallinales) and the other dominated by fleshy algae, in an area that has been exposed to important levels of anthropogenic disturbance, mainly pollution (including changed sedimentation regimes) in the recent past (bay of Marseilles, France). In comparison with less strongly impacted Mediterranean regions, the macrophyte assemblages in the bay of Marseilles were characteristic in terms of species identity and richness of coastal detritic macrophyte assemblages. However, extremely low species abundance (cover) was observed. As far as invasive species were concerned, Caulerpa racemosa var. cylindracea was only abundant in the rhodolith assemblage whereas the two invasive Rhodophyta Asparagopsis armata and Womersleyella setacea were mainly found in the fleshy algae assemblage. The seasonality observed in the Rhodolith assemblage seemed to be related to the development of C. racemosa var. cylindracea and did not follow the typical pattern of other Mediterranean assemblages. This study represents the first study of coastal detritic assemblages invaded by C. racemosa var. cylindracea.     

Hard coastal-defence structures as habitats for native and exotic rocky-bottom species

The use of hard coastal-defence structures, like breakwaters and seawalls, is rapidly increasing to prevent coastal erosion. We compared low-shore assemblages between wave-protected and wave-exposed habitats on breakwaters along a sandy shore of Tuscany (North-Western Mediterranean). Assemblages were generally characterized by a low diversity of taxa, with space monopolized by Mytilus galloprovincialis and Corallina elongata on the seaward side of breakwaters and by filamentous algae on the landward side. Assemblages in wave-protected habitats were characterized by greater temporal stability than those in exposed habitats and supported non-indigenous macroalgae such as Caulerpa racemosa and Codium fragile ssp. tomentosoides. Hence, the introduction of hard coastal-defence structures in otherwise soft-bottom dominated areas, attracting native and exotic rocky-bottom species, should be of great concern for the conservation of marine biodiversity at local and regional scales and for the management of biological invasions.     

Comparison between amphipod assemblages associated with Caulerpa racemosa var. cylindracea and those of other Mediterranean habitats on soft substrate

The spread of the invasive alga Caulerpa racemosa var. cylindracea in shallow-water habitats can present different faunal assemblage composition. We compared the amphipod assemblages associated with C. racemosa and natural habitats found on shallow-water Mediterranean soft substrata. Four vegetated habitats were compared: C. racemosa, Caulerpa prolifera, Cymodocea nodosa and Posidonia oceanica with unvegetated substrata. Samples were collected during two sampling periods (September 2004 and March 2005). A total of 63 amphipod species were recorded. The results showed that the vegetated habitats sampled, including C. racemosa stands, supported a higher abundance and species richness of amphipods. Furthermore, the assemblage structure differed between the different habitats, while the abundance of some species was significantly different, depending on habitat. For example, Microdeutopus obtusatus was favoured by C. racemosa habitat; Ampelisca diadema was associated with C. prolifera beds; and Hyale schmidti was more abundant in P. oceanica meadows. Habitat invasion by C. racemosa can exert an important influence on biotic assemblages, modifying habitat structure and associated fauna.     

Functional changes due to invasive species: Food web shifts at shallow Posidonia oceanica seagrass beds colonized by the alien macroalga Caulerpa racemosa

Multiple stable isotope analyses were used to examine the trophic shifts at faunal assemblages within the invading macroalga Caulerpa racemosa in comparison to established communities of Posidonia oceanica seagrass meadows. Sampling of macrobenthic invertebrates and their potential food sources of algal mats and seagrass meadows in Mallorca (NW Mediterranean) showed differences in species composition of faunal and primary producers among seagrass and C. racemosa. Accordingly, changes in food web structure and trophic guilds were observed, not only at species level but also at community level. The carbon and nitrogen isotope signatures of herbivores, detritivores and deposit feeders confirmed that the seagrass provided a small contribution to the macrofaunal organisms. δ¹³C at the P. oceanica seagrass and at the C. racemosa assemblages differed, ranging from −6.19 to −21.20‰ and −2.67 to −31.41‰, respectively. δ¹⁵N at the Caulerpa mats was lower (ranging from 2.64 to 10.45‰) than that at the seagrass meadows (3.51–12. 94‰). Significant differences in isotopic signatures and trophic level among trophic guilds at P. oceanica and C. racemosa were found. N fractionation at trophic guild level considerable differed between seagrass and macroalgae mats, especially for detritivores, deposit feeders, and herbivores. Filter feeders slightly differed with a relatively lower N signal at the seagrass and CR values at community level and at trophic guild level were higher in the C. racemosa invaded habitats indicating an increase in diversity of basal resource pools. C. racemosa did seem to broaden the niche diversity of the P. oceanica meadows it colonised at the base of the food web, may be due to the establishment of a new basal resource. The extent of the effects of invasive species on ecosystem functioning is a fundamental issue in conservation ecology. The observed changes in invertebrate and macrophytic composition, stable isotope signatures of concomitant species and consequent trophic guild and niche breadth shifts at invaded Caulerpa beds increase our understanding of the seagrass systems.     

Effects of the spread of the alga Caulerpa racemosa var. cylindracea on the sponge assemblage from coralligenous concretions of the Apulian coast (Ionian Sea, Italy)

The present work investigated the modifications induced by the spread of the green macroalga Caulerpa racemosa var. cylindracea (Bryopsidales, Chlorophyta) on the sponge assemblage of Apulian coralligenous concretions (Ionian Sea – Torre Ovo, Italy). The study of qualitative and quantitative sponge composition was carried out before (2004) and after (2006) the spread of this invasive alga by means of traditional (quadrat scraping) and photographic sampling methods. Results indicate that the spread of the green alga is concomitant with a significant decrease in percentage sponge cover both on horizontal- and on vertical-oriented substrates. In addition, strong modifications to the structure of the community in terms of repartition of the available substrate have been observed since the algal spread. Conversely, no major changes have affected the specific composition of the sponge assemblage, suggesting that at this stage of colonization the algal spread has not produced a loss of sponge biodiversity. However, there is a clear need to monitor closely the C. racemosa invasion to verify its long-term impact on the sponge assemblage.     

Assessment of substratum effect on the distribution of two invasive Caulerpa (Chlorophyta) species

Two-year monitoring of the invasive marine Chlorophyta Caulerpa taxifolia and Caulerpa racemosa var. cylindracea shows the great influence of substratum on their spatial distribution. The cover of C. taxifolia and C. racemosa was measured in shallow (<8 m) areas indicating that these species are more abundant in rocks with photophilic algae and in the dead matte of the seagrass Posidonia oceanica than in sand or inside the P. oceanica meadow. A short-term experiment comparing the persistence of C. taxifolia and C. racemosa planted either in a model of dead matte of P. oceanica or in sand shows that the persistence of these species was higher in the dead matte model than in sand. Correlative evidence suggests that C. taxifolia and C. racemosa tolerate near-bottom orbital velocities below 15 cm s^?1 and that C. taxifolia cover declines at velocities above that value. These results contribute to understand the process of invasion of these Caulerpa species predicting which substrata would be more susceptible to be invaded and to the adoption of appropriate management strategies.     

Limited impact of Sargassum muticum on native algal assemblages from rocky intertidal shores

Invasive species represent a serious threat to natural ecosystems through a range of negative effects on native species in the region invaded. The invasive species Sargassum muticum has invaded several temperate regions worldwide including the Galician rocky shoreline (northwestern Spain) in Western Europe. The main aim of this study was to assess if colonization by S. muticum has any effect on native algal assemblages by experimental removal of S. muticum. We predicted that in those plots where S. muticum plants were removed, the structure of native algal assemblages would differ from that in plots where S. muticum plants were untouched. In addition, we predicted that the effect of Sargassum removal would be more important than other causes of variability at the small scale investigated. Results indicated limited impact of S. muticum on native assemblages. The impact was only evident on the total number of native taxa and two understory morpho-functional groups, filamentous and foliose algae, rather than on the entire macroalgal assemblages.     

Temporal and spatial variability in shallow- and deep-water populations of the invasive Caulerpa racemosa var. cylindracea in the Western Mediterranean

The invasive green alga Caulerpa racemosa var. cylindracea represents an important threat to the diversity of Mediterranean benthic coastal ecosystems by interfering with native species and modifying benthic assemblages. The present study deals, for the first time, with the temporal and spatial variability of the biomass and phenology of C. racemosa considering both deep- and shallow-water populations. Two sampling depths (30 and 10 m) were sampled at three different rocky bottom sites every 3 months in the Archipelago of Cabrera National Park (Western Mediterranean). All morphometric variables analysed showed a spatial variation and temporal patterns depending on depth. Between depths, C. racemosa biomass, stolon length, number of fronds and frond length were usually significantly higher at deep-water populations, suggesting that C. racemosa grows better in deep-waters. Deep- and shallow-water populations displayed a high temporal variation although no evidence of seasonal patterns was found, in contrast with what has been reported by other authors. The sources of this variability are still unknown but probably both physical factors and differential herbivory pressures display a key role.     

Settlement and colonization of epi‐endobenthic communities on calcareous substrata in an underwater archaeological site

Underwater archaeological sites represent assets of great value that are subjected to physical and biological stresses. The study of biological settlement on calcareous substrata is the first step to understanding the biodeterioration processes that take place on man‐made structures (marble, statues, columns and other architectonic elements) in marine environments. To accomplish this objective, calcareous panels (30 × 30 × 2 cm) were immersed for 3, 6, 9 and 12 months along a north?south transect in marine protected areas of Baia (Naples). Both epilithic and endolithic assemblages were analysed: the latter was observed by scanning electron microscope after the cast‐embedding technique had been used. After 1 year the epilithic community was still changing, with algae and pioneer species overgrown by encrusting organisms, and the endolithic assemblage dominated by cyanobacteria/chlorophyte traces. Changes of epilithics lead changes on endolithics. The initial phototrophic borers were replaced by more sciophilous and competitive species; moreover, the first macroboring organisms arrived after the settlement of grazers. After 12 months of immersion, the rate of bioerosion was high, the material was deeply bored and all of the limestone surface was heavily covered.     

西沙群岛海产绿藻的研究Ⅲ

本文继续报道产于我国广东省西沙群岛的海产绿藻9种,除2种已有记载外,其余7种都是西沙群岛的新记录,其中1种为新种一簇囊缢丝藻Boodleopsis aggregata sp.nov.,2种为我国的新记录—太坦松藻Codium taitense Setchell,钱币状蕨藻Caulerpa nummularia (Harv. ) Reinke。     

Variability in the structure of epiphytic assemblages of the Mediterranean seagrass Posidonia oceanica in relation to depth

The aim of the present study was to evaluate whether the variability in the structure of the epiphytic assemblages of leaves and rhizomes of the Mediterranean seagrass Posidonia oceanica differed between depths at a large spatial scale. A hierarchical sampling design was used to compare epiphytic assemblages at two different depths (10 and 20 m) in terms of both species composition and abundance and patterns of spatial variability in the Tuscan Archipelago (North Western Mediterranean Sea, Italy). Results showed significant differences in the structure of assemblages on rhizomes and leaves at different depths. These differences were related to species composition and abundance; differences were not significant for total biomass, total percentage cover and percentage cover of animals and algae. Whereas the higher variability was observed among shoots in all the studied systems, patterns of spatial variability at the other spatial scales investigated differed between the two studied depths. Moreover, in the present study, analogous patterns between depths resulted for both the assemblages of leaves and rhizomes, suggesting that factors that change with depth can be responsible for the spatial variability of both the assemblages (leaves and rhizomes), and operate regardless of the microclimatic conditions and the structure of assemblages.     

Algae as hosts for epifaunal bryozoans: Role of functional groups and taxonomic relatedness

Macroalgae build biogenic habitats which give shelter and provide a suitable physical environment for a great variety of organisms. Structural complexity of algal substrates may influence the composition of their attached epifauna. The aim of this study is to test whether the taxonomic relatedness of the algal hosts and the functional groups to which they belong influence the species richness and composition of their epifaunal bryozoans. We analysed 36 algal genera from the Atlantic coast of South America between 42°S and Cape Horn. Changes in bryozoan species richness (number of species) among different algal functional groups (filamentous algae, foliose algae, corticated foliose algae, corticated macrophytes) were non-significant. The composition of the epifaunal assemblages differed significantly only between filamentous and foliose algae. Sheet-like bryozoans (i.e. encrusting, pluriserial colonies) were more frequent on foliose than on filamentous algae, while runner-like species (i.e. uniserial stolons) were characteristic epibionts on filamentous thallii. Similarity of bryozoan assemblages increased with increasing taxonomic relatedness of their hosts. As most filamentous seaweeds analysed in this study are members of the Order Ceramiales, the influence of algal taxonomic relatedness and functional groups on the composition of their bryozoan assemblages can be viewed as two different aspects of the same phenomenon.     

Seasonal variation and structure of a decapod (Crustacea) assemblage living in a Caulerpa prolifera meadow in Cádiz Bay (SW Spain)

The decapod taxocoenosis living in shallow muddy bottoms with the green algae Caulerpa prolifera was studied monthly between February 1994 and January 1996 in the Inner Bay of Cádiz (SW Spain). More than 32,000 specimens belonging to 35 species were collected. Six species were dominant (representing the 85.8% of the total number of specimens), but the structure of the taxocoenosis was regulated by the Hippolyte species, Sicyonia carinata, Palaemon adspersus and Liocarcinus arcuatus. There was no significant qualitative difference between years. There was no clear change in the dominance of groups of species during the year, as happened in the outer Bay. This is probably due to the sheltered character of the area and the more stable and dense vegetal cover, but some seasonal differences were found.The benthic characteristics of the Inner Bay of Cádiz, such as shallow soft bottoms of fine and muddy sediments and the presence of macrophytes (seagrasses and seaweeds) might be key factors influencing the composition and structure of the general and seasonal decapod assemblage. In spite of human impacts on the bay (e.g. aquaculture activities, sewage), the values of the diversity, equitability and richness indexes appeared stable over time (higher than those found in outer adjacent areas) and no significant differences between years were found.     

Spatial variability of the surf zone fish and macroinvertebrate community within dissipative sandy beaches in Oregon,USA

Habitat heterogeneity can influence biological communities by providing a diversity of areas that can be occupied by different species. Sandy beach surf zones are often considered homogenous environments; however, sand bars moved by currents and waves can produce trench‐like shapes or troughs that provide heterogeneity. The influence of habitat heterogeneity produced by sand movement is unclear despite the fact that surf zones are an important habitat for larval and juvenile fish and macrocrustaceans. To determine if, and how, the fish and macroinvertebrate communities present in trough and non‐trough or flat areas of Oregon surf zones differ, we compared species assemblages in both areas at three beaches adjacent to estuary mouths over 2 years. Troughs had different communities compared with flat areas, with higher total catch (mean ± SD = 123.2 ± 122.1 versus 43.6 ± 44.5 individuals × 100 m^?2) and taxon richness (6.7 ± 2.7 versus 4.0 ± 2.3 taxa); these differences were potentially due to water movement, prey availability and sediment size. The fish and macroinvertebrate communities did not vary between years but there were significant differences among beaches, with the most distinct community present at the only beach adjacent to an estuary without a jetty at its mouth, which was possibly due to higher species movement between the surf zones and estuary. Fish and macrocrustacean surf zone communities varied spatially within and among beaches in relation to habitat heterogeneity provided by sand movement and, potentially, the influence of adjacent habitats.     

Expansion of the macroalga Caulerpa racemosa and changes in softbottom macrofaunal assemblages in Moni Bay,Cyprus

The recent expansion of the Red Sea macroalga Caulerpa racemosa and its impact on the diversity and abundance of macrobenthos were examined and compared in the summers of 1992 and 1997, in Moni Bay, Cyprus. The phytobenthic community of the bay in 1992 was dominated by the seagrass Posidonia oceanica while, in 1997, the Lessepsian migrant C. racemosa became the most dominant, forming extensive mats. Changes in the vegetation system in Moni Bay have caused significant compositional changes in macrofaunal assemblages. A total of 178 individuals of 62 species are recorded. The composition of the macrofauna in 1992 was dominated by gastropods (44 %), crustaceans (22%), bivalves (17 %), polychaetes (11 %) and echinoderms (6 %). In 1997, the gastropods and crustaceans had decreased to 13 % and 16 % respectively, while, polychaetes had increased to 38 % becoming the most dominant taxon. Bivalves and echinoderms also increased to 22 % and 11 %, respectively, in 1997. The proliferative growth of C. racemosa imposed successional changes on the macrofaunal assemblages in Moni Bay, Cyprus, between 1992 and 1997. It remains to be tested whether the expansion of C. racemosa is related to the increase of water temperature associated with global warming or nutrient inputs or with the differences in the life history characteristics of this migrant vs. native algal species.