Effects of macroalgal identity on epifaunal assemblages: native species versus the invasive species Sargassum muticum

Seaweeds are a refuge from stressful conditions associated with life on rocky intertidal shores, and there is evidence that different macrophytes support different assemblages of mobile epifauna. Introduction of non-indigenous macroalgae may have a great impact on associated epifaunal assemblages and ecosystem processes in coastal areas. Previous studies have reported conflicting evidences for the ability of epifauna to colonize non-indigenous species. Here, we analyzed epifaunal assemblages associated with three species of macroalgae that are very abundant on intertidal shores along the Galician coast: the two native species Bifurcaria bifurcata and Saccorhiza polyschides and the invasive species Sargassum muticum. We collected samples of each species from three different sites at three different times to test whether variability of epifaunal assemblages was consistent over space and time. Epifaunal assemblages differed between the three macroalgae. Results suggested that stability and morphology of habitat played an important role in shaping the structure of epifaunal assemblages. This study also showed that the invasive S. muticum offered a suitable habitat for many invertebrates.     

Diurnal, horizontal and vertical dispersal of kelp-associated fauna

The kelp Laminaria hyperborea is host for a rich fauna of mobile invertebrates. Dispersal patterns of these animals may be crucial for their availability to visual predators like fish, which are known to search for food in these productive habitats. Diurnal, horizontal and vertical dispersal within and out of the kelp forest were studied by analysing colonization of artificial substrata placed among kelps. The species composition of the fauna was different on three parts of the kelp: lamina, stipes (with epiphytes) and holdfast. The artificial substrata were colonized mainly by species associated with kelp. More species and individuals colonized the artificial substrata at night than during the day. The most abundant faunal groups on the kelps and the artificial substrata were amphipods and gastropods. Both groups dispersed at a significantly higher rate at night than during the day. Rapid horizontal dispersal out of the kelp forest was found. The artificial substrata were also colonized quickly by kelp fauna at all vertical levels inside and above the kelp forest. However, species associated with the kelp holdfast tended to disperse close to the bottom, while stipe-associated fauna moved through all parts of the kelp forest and even above the canopy layer. A high dispersal rate appears common amongst the mobile species living on kelp and seems to be advantageous, despite the risks involved in emigrating from the habitat and being exposed to predators. Higher activity at night may reduce availability to predators.     

Seasonal variation in epifaunal communities associated with giant kelp (Macrocystis pyrifera) at an upwelling‐dominated site

Kelp forests are highly productive and species‐rich benthic ecosystems in temperate regions that provide biogenic habitat for numerous associated species. Diverse epifaunal communities inhabit kelp sporophytes and are subject to variations in the physical environment and to changes experienced by the kelp habitat itself. We assessed seasonal variations in epifaunal invertebrate communities inhabiting giant kelps, Macrocystis pyrifera, and their effects on this seaweed. Six seasonal samplings were conducted over a year at an upwelling‐dominated site in northern‐central Chile where physical conditions are known to fluctuate temporally. More than 30 taxa were identified, among which peracarid crustaceans stood out in both diversity and abundance. Species richness and abundance differed among sporophyte sections (holdfast and fronds) and throughout the year. The frond community was dominated by two grazers (the amphipod Peramphithoe femorata and the isopod Amphoroidea typa), while suspension feeders, grazers, and omnivores (the amphipod Aora typica, the isopod Limnoria quadripunctata, and polychaetes) dominated the holdfasts. Abundances of the dominant species fluctuated throughout the year but patterns of variation differed among species. The most abundant grazer (P. femorata) had highest densities in summer, while the less abundant grazer (A. typa) reached its peak densities in winter. Interestingly, the area of kelp damaged by grazers was highest in autumn and early winter, suggesting that grazing impacts accumulate during periods of low kelp growth, which can thus be considered as ‘vestiges of herbivory past.’ Among the factors determining the observed seasonal patterns, strong variability of environmental conditions, reproductive cycles of associated fauna, and predation by fishes vary in importance. Our results suggest that during spring and early summer, bottom‐up processes shape the community structure of organisms inhabiting large perennial seaweeds, whereas during late summer and autumn, top‐down processes are more important.     

Disentangling the links between habitat complexity and biodiversity in a kelp‐dominated subantarctic community

Habitat complexity is one of the most important factors modulating species diversity. This feature comprises several interrelated attributes, such as number, size, and spatial arrangement of complexity‐forming elements. However, the separate and joint effects of these attributes on diversity and community structure are still not well understood. Here, we assess the relationships between several structural‐complexity attributes of the subantarctic kelp Lessonia flavicans and species richness, total abundance, and structure of kelp‐associated macrobenthic communities. We predicted that longer thalli and larger holdfasts favor greater species richness and total abundance of invertebrate organisms. To test the prediction, an observational sampling program was established in two sites of the Strait of Magellan. Uni‐ and multivariate analyses revealed both positive and negative effects of kelp structural‐complexity attributes on diversity. Holdfast diameter and maximum frond length, followed by thallus wet weight, had the strongest positive fits to species richness and total abundance; the number of stipes, on the other hand, was negatively associated with both response variables. Longer fronds were associated with greater abundances of spirorbid polychaetes. Larger holdfasts supported larger abundances of Nereididae and Terebelidae polychaetes and the limpet Nacella mytilina. Contrarily, kelps with longer fronds and more stipes supported fewer amphipods. In this way, we demonstrate that different dimensions of habitat complexity can have contrasting effects on diversity and community structure, highlighting the fundamental role of multiple dimensions of kelp habitat complexity for local biodiversity.     

Nonlinear multivariate models of successional change in community structure using the von Bertalanffy curve

von Bertalanffy curves were used to describe the nonlinear relationship between assemblages inhabiting holdfasts of the kelp Ecklonia radiata and the volume of the holdfast. This was done using nonlinear canonical analyses of principal coordinates (NCAP). The volume of the holdfast is a proxy for the age of the plant and, thus, the canonical axis is a proxy for succession in the marine invertebrate community inhabiting the holdfast. Analyses were done at several different taxonomic resolutions on the basis of various dissimilarity measures. Assemblages in relatively large holdfasts demonstrated ongoing variation in community structure with increasing volume when the dissimilarity used was independent of sample size. Smaller holdfasts had proportionately greater abundances of ophiuroids and encrusting organisms (bryozoans, sponges, ascidians), while larger holdfasts were characterised by proportionately greater abundances of crustaceans, polychaetes and molluscs. Such linear and nonlinear multivariate models may be applied to analyse system-level responses to the growth of many habitat-forming organisms, such as sponges, coral reefs, coralline algal turf or forest canopies.     

Regional-scale analysis of subtidal rocky shore community

The French monitoring network, REseau BENThique (REBENT), was launched by the Ministry of the Environment in 2003 following the 1999 Erika oil spill. REBENT aimed to acquire baseline knowledge of coastal benthic habitat distributions with a special focus on biological diversity. This study analyzed data from 38 subtidal rocky reef sites collected by a single diving team of marine biologists along the coast of Brittany from 2004 to 2010. At each site, the depth limits of the algal belts were determined between 0 and ?40 m Chart Datum (CD); the flora and fauna compositions and abundances were sampled at ?3 and ?8 m CD. A total of 364 taxa (156 flora and 208 fauna), belonging to 12 phyla, were identified. The results showed that the depth limit and density of kelp beds increased as water turbidity decreased; moreover, several changes in community structure could be related to water turbidity and temperature. Thus, northern and southern Brittany showed strong differences in diversity and structure of the dominant kelp species (Laminaria hyperborea and Saccorhiza polyschides). The results from this kelp habitat composition survey (dominant kelp species and indicator species) provided important information for local pressure assessments, like increases in turbidity. The data also provided a reference that could be useful for detecting changes in coastal water temperatures due to global warming.     

Short-term dispersal of kelp fauna to cleared (kelp-harvested) areas

The kelp Laminaria hyperborea forms large forests and houses a numerous and diverse fauna, especially in the kelp holdfast and stipe epiphytes. Kelp harvesting creates cleared areas and fragmentizes the kelp forest. We investigated the dispersal ability of kelp fauna to cleared, harvested areas by studying their colonization pattern to artificial substrata (kelp mimics) exposed for a short (3 days) and longer time period (35 days) at different sites within the kelp forest (one site) and at a cleared area (two sites). Most of the kelp fauna (111 species) showed a rapid dispersal and colonized the artificial substrata within the cleared area. The similarity of the faunal community in the mimics with the natural kelp holdfast community increased with the length of the exposure period. During the experiments, 87% of the mobile species in the kelp plants were found in the kelp mimics, indicating good dispersal for slow-moving animals like gastropods, polychaetes and tube-building crustaceans. Relating the frequency of the different faunal groups in the untrawled kelp forest to their frequency in the kelp mimics, showed gastropods, amphipods and decapods to have relatively high dispersal rates, whereas isopods, bivalves, polychaetes and tanaids showed a lower dispersal rate than expected. Amphipods dispersed as juveniles and adults. No significant differences were found between the faunal composition and number of species in the mimics placed inside the kelp forest and in the cleared area. Remaining holdfasts and pebbles were identified as refuges/alternative habitats in the harvested area, and may together with the nearest kelp vegetation, serve as sources for colonization to new substrata. The high dispersal ability of most of the kelp fauna provides maintenance of the faunal composition of disturbed habitats and ensures colonization of recovering algal habitats regardless of reproduction strategy.     

Polychaete fauna associated with holdfasts of the large brown alga Himantothallus grandifolius in Admiralty Bay, King George Island, Antarctic

The polychaete community associated with holdfasts of the brown alga Himantothallus grandifolius in Admiralty Bay has been studied. It is the first study of its kind in this area and only the second in the Antarctic. Samples were collected in the summer season of 1979/1980 from a depth range of 10–75 m. Seventy-eight species were found on 19 holdfasts. The community was dominated by Brania rhopalophora and Neanthes kerguelensis. Analysis of similarity showed that polychaete fauna associated with this habitat did not show any partitioning related to depth. Regression analysis showed that densities of both species and individuals decreased with increased holdfast volume. A positive correlation was found between the number of individuals and holdfast volume. Polychaetes from 10 feeding guilds were found with dominance of macrophagous motile herbivores and sessile filter feeders. The complex habitat provided by holdfasts is a shelter for a rich polychaete fauna and may function as important protection from disturbance in the shallow areas of Admiralty Bay.     

Seasonal changes in taxon richness and abundance of mobile invertebrates inhabiting holdfast of annual kelp Ecklonia radicosa (Phaeophyceae,Lessoniaceae) at the central Pacific coast of Japan

To investigate seasonal changes in the taxon richness and abundance of mobile invertebrates inhabiting holdfasts of the warm temperate annual kelp Ecklonia radicosa, five holdfasts were collected monthly at the central Pacific coast of Japan from April to November 2014. During the study period, there was little variation in holdfast height and diameter, which ranged from 5.9 to 8.5 cm and from 7.1 to 10.8 cm, respectively. In total 7087 animals were collected from 40 holdfasts (177.2 individuals inds./holdfast, on average). The number of mobile invertebrates gradually increased from May (15 ± 9.9 inds./holdfast) to August (346 ± 152.5 inds./holdfast), with over 300 inds./holdfast until October before rapidly decreasing in November (110 ± 85.6 inds./holdfast). Similarly, taxon richness increased gradually from April (4.3 ± 1.0 taxa/holdfast) to August (11.0 ± 3.7 taxa/holdfast), and decreased in November (8.6 ± 2.3 taxa/holdfast). Interestingly, hundreds of mobile invertebrates inhabited holdfasts of kelp plants that had shed their blade in October and November. Taxon composition from August to October and the number of invertebrates from July to November were comparable to data previously reported for perennial kelps. In this study, the importance of annual Ecklonian species as biogenic habitats was demonstrated for the first time.     

The kelp fauna of northeast Britain. II. Multivariate classification: Turbidity as an ecological factor

Detailed faunal data relating to a sublittoral survey of kelp holdfasts (Laminaria hyperborea (Gunn.) Fosl.) in northeast Britain are presented. These data have been ordered by multivariate statistical techniques. Analytical strategy-is outlined and results pertaining to the classificatory techniques of association analysis (ASOC), predictive attribute analysis (PRED) and divisive information analysis (DINF) using qualitative data are given. Normal and inverse hierarchies wree computed using ASOC and DINF and used to construct two-way tables. Recognition of environmentally definie species assemblages is possible from a consideration of particular analyses. Taking into account certain methodological and sampling limitations, a definition of groups of ubiquitous, clear-water, turbidity indifferent and turbid water species may be recognized; the significant species for which purposes are only one-third of the total number considered.A primary correlation between the local composition of the holdfast fauna and turbidity is indicated. Subsidiary pollution effects cannot be ruled out but their definition becomes complicated by the intervention at lower levels of heterogeneity of other correlating factors, e.g., holdfast morphology. The danger of attributing causal relationships to statistical correlations is stressed throughout.     

Climate‐driven substitution of habitat‐forming species leads to reduced biodiversity within a temperate marine community

Aim

In marine ecosystems, habitat‐forming species (HFS) such as reef‐building corals and canopy‐forming macroalgae alter local environmental conditions and can promote biodiversity by providing biogenic living space for a vast array of associated organisms. We examined community‐level impacts of observed climate‐driven shifts in the relative abundances of two superficially similar HFS, the warm‐water kelp Laminaria ochroleuca and the cool‐water kelp Laminaria hyperborea.

Location

Western English Channel, north‐east Atlantic

Methods

We compared algal and invertebrate assemblages associated with kelp stipes and holdfasts, across multiple sites and sampling events. Significant differences were recorded in the structure of assemblages between the host kelp species at each site and event.

Results

Assemblages associated with stipes of the cool‐water HFS were, on average, >12 times more diverse and supported >3600 times more biomass compared with the warm‐water HFS. Holdfast assemblages also differed significantly between species, although to a lesser extent than those associated with stipes. Overall, assemblages associated with the warm‐water HFS were markedly impoverished and comprised far fewer rare or unique taxa.

Main conclusions

While previous research has shown how climate‐driven loss of HFS can cause biodiversity loss, our study demonstrates that climate‐driven substitutions of HFS can also lead to impoverished assemblages. The indirect effects of climate change remain poorly resolved, but shifts in the distributions and abundances of HFS may invoke widespread ecological change, especially in marine ecosystems where facilitative interactions are particularly strong.     

Consistency and variation in kelp holdfast assemblages: Spatial patterns of biodiversity for the major phyla at different taxonomic resolutions

The focus of this study was to measure natural spatial variability in the biodiversity of fauna inhabiting kelp holdfasts in northeastern New Zealand at several spatial scales: from meters up to hundreds of kilometers. We wished to test the hypothesis that multivariate variation and biodiversity would vary significantly at different spatial scales in different ways for the major phyla in the holdfast community (Arthropoda, Annelida, Mollusca and Bryozoa). Biodiversity was considered in terms of richness, total abundance, structural composition (as measured by the Bray-Curtis dissimilarity measure) and taxonomic breadth for each major phylum and for the assemblage as a whole. We also examined the effect of taxonomic resolution on multivariate patterns. Species richness and total abundance increased with increases in holdfast volume. Multivariate variation was greatest at the smallest spatial scale for all phyla, but different phyla showed different patterns of multivariate variation at different spatial scales. Variations among locations at the largest spatial scale were primarily due to differences in the composition and richness of bryozoans and molluscs. Location effects became less and less distinct with decreases in taxonomic resolution. There were very few significant differences in richness or abundance for holdfasts of a given volume, taxonomic breadth did not vary significantly across locations, nor did the proportional abundances of phyla. These consistencies across large spatial scales in the absence of environmental impacts and results from other studies suggest that holdfast communities in New Zealand systems would provide a useful model assemblage against which future impacts may be detected as changes in proportions of component phyla. In addition, high variability detected at small and large scales at the species level, especially for bryozoans and molluscs, suggest that these communities may also provide unique opportunities for studying and understanding sources and functions of marine biodiversity.     

Multivariate dispersion as a measure of beta diversity

Beta diversity can be defined as the variability in species composition among sampling units for a given area. We propose that it can be measured as the average dissimilarity from individual observation units to their group centroid in multivariate space, using an appropriate dissimilarity measure. Differences in beta diversity among different areas or groups of samples can be tested using this approach. The choice of transformation and dissimilarity measure has important consequences for interpreting results. For kelp holdfast assemblages from New Zealand, variation in species composition was greater in smaller holdfasts, while variation in relative abundances was greater in larger holdasts. Variation in community structure of Norwegian continental shelf macrobenthic fauna increased with increases in environmental heterogeneity, regardless of the measure used. We propose a new dissimilarity measure which allows the relative weight placed on changes in composition vs. abundance to be specified explicitly.     

Evaluating stress in rocky shore and shallow reef habitats using the macrofauna of kelp holdfasts

Kelp often dominates hard rocky substrata in the coastal watersof temperate, boreal and subantarctic regions of the world's oceans. The habitat providedby kelp holdfasts supports high species richness and high abundance of a range of invertebratetaxa. Studies conducted on the effects of stress on these communities have indicated that theyare sensitive to stressors and that the response is often predictable. With the growing body ofliterature on natural variation in holdfast community structure, studies of the impacts of stressare becoming more readily interpretable making the macrofauna of this habitat a potentiallyimportant tool for evaluating anthropogenic effects on hard substrata in coastal environments. Limitations of the use of holdfast fauna in studies of stress include the fact that brownseaweeds may disappear from polluted areas of reef, and where they persist, holdfaststructure may be altered directly by the stressor. An additional problem is that manipulative,experimental studies are difficult. Artificial holdfasts may provide a means by which thedemonstrated sensitivity of the holdfast community can be combined with a manipulative approach to testspecific hypotheses related to effects of stress in shallow coastal environments.     

Ecological effects of harvesting Lessonia (Laminariales,Phaeophyta) in central Chile

Lessonia nigrescens and L. trabeculata are economically important canopy-forming kelps in Chile. Experimental harvesting of stipes above the first dichotomy reduces stipe movement and inter-stipe friction, allowing the development of a heavy epiphytic load and increased grazing. Complete stipe removal leads to holdfast death as neither species is able to simultaneously regenerate all stipes. The invertebrate fauna inside the holdfast does not respond to upper canopy changes, but mortality does occur in partial or complete plant removals. Kelp removal also affects inter-plant distances, results in increased access of grazers to the outside and inside of kelp holdfasts, reduces recruitment of other algal species, and modifies the morphology of L. trabeculata such that the plants become more susceptible to removal by water movement.     

Does restoration of a habitat‐forming seaweed restore associated faunal diversity?

Declines of habitat‐forming organisms in terrestrial and marine systems can lead to changes in community‐wide biodiversity. The dominant habitat‐forming macroalga Phyllospora comosa (Fucales) went locally extinct along the metropolitan coastline of Sydney in the 1980s. However, the consequences of that disappearance to the associated faunal diversity in these habitats, and whether Phyllospora is ecologically redundant with respect to the biodiversity it supports, are not known. Efforts are underway to restore Phyllospora, and the capacity to enhance local biodiversity is an important component of the rationale for restoration. We compared epifaunal diversity (abundances and composition) between Phyllospora and two other co‐occurring habitat‐forming algae, the kelp Ecklonia radiata and the fucoid Sargassum vestitum, and determined whether Phyllospora transplanted to Sydney developed different epifaunal communities than undisturbed thalli and controls. Where the 3 species naturally co‐occurred, Phyllospora supported different abundances of taxa than Ecklonia and Sargassum, as well as different composition at finer scales, which suggests that this species is not completely redundant and that its disappearance may have affected local biodiversity. Similarly, assemblages on transplanted Phyllospora differed from those on Ecklonia and Sargassum at restored sites, but did not always resemble assemblages from extant natural Phyllospora populations, even 18 months after transplantation. These experiments indicate that restoration of key habitat‐forming seaweeds not only recovers the algal species but also reduces risks of losing habitat diversity for epifauna and their consumers. However, restoration of all the original biodiversity associated with these seaweeds can be a difficult, complex, and long‐term process.     

Spatial and temporal variability of the benthic assemblages associated to the holdfasts of the kelp <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> in the Straits of Magellan,Chile

Macrocystis pyrifera (L.) C. Agardh is a characteristic macroalga in the Magellan region covering almost 30% of the shallow coastal waters. The focus of this study was to analyse the spatial and seasonal patterns in macrofauna communities associated to the holdfasts of Macrocystis pyrifera at two study sites in the Straits of Magellan, South Chile. In total, 114 species from 10 major taxa were isolated from the holdfasts. MDS clearly separated the holdfast fauna collected in different seasons, with autumn and winter collections being richer in terms of species richness and abundance as compared to the spring and summer situation. MDS also clearly separated the holdfast associated faunas of the two study sites, Bahía Laredo and Fuerte Bulnes. The community structure and species composition of the associated macro-invertebrates and vertebrates appeared rather heterogeneous, probably due to the extremely heterogeneous environmental conditions along the entire coastline of the Subantarctic Magellan region.     

Biodiversity patterns of macro-epifauna on different lamina parts of Laminaria digitata and Saccharina latissima collected during spring and summer 2004 in Kongsfjorden, Svalbard

The epifauna associated with two of the most common species of kelp in the Arctic, Laminaria digitata (Hudson) Lamouroux, 1813 and Saccharina latissima (Linnaeus) Lane, , Mayes, Druehl and Saunders 2006 [synonym: L. saccharina (Linnaeus) Lamouroux, 1813] were examined in Kongsfjorden, Svalbard. The aim of this study was to test whether species richness of epifauna was influenced by seasonality of recruitment supply (spring and summer) or by age of the substrate (lamina). From 20 algae specimens (ten—L. digitata, ten—S. latissima), a total of 27 epifaunal taxa were identified with bryozoans as the most species-rich group. Total species richness ranged from one to nine species per lamina. No significant difference in species richness was observed between the two kelp species. In both macroalgae, more epifauna taxa were registered on lamina in May (mean 6) than in August (mean 3). This indicates that the time chosen for sampling had a significant impact on the species richness pattern. The number of epifaunal species was significantly negatively correlated with increasing age of the lamina. There are indications that diversity of the epifauna on lamina is influenced to a large extent by substrate (lamina) surface area, environmental stress and the presence of meroplankton in the water column.