Patterns of abundance and assemblage structure of epifauna inhabiting two morphologically different kelp holdfasts

The mobile fauna associated with two sympatric kelp species with different holdfast morphology (Saccorhiza polyschides and Laminaria hyperborea) was compared to test for differences in the assemblage structure of holdfast-associated mobile epifauna. A total of 24,140 epifaunal individuals were counted from 30 holdfasts of each kelp species. Overall epifaunal abundances exceeded faunal abundances previously reported from holdfasts of other kelps. Three taxonomic groups, Amphipoda, Mollusca, and Polychaeta, accounted for ca. 85% of all individuals. Total abundances increased with the amount of habitat available, quantified either as the volume or the area provided by the holdfasts. The multivariate structure of the epifaunal assemblage did not differ between holdfasts of the two kelp species. However, epifaunal assemblages responded differentially to the habitat attributes provided by each type of kelp holdfast: multivariate variation in the assemblage structure of epifauna was mostly explained by holdfast area and volume for L. hyperborea, and by the surface-to-volume ratio for S. polyschides holdfasts. Therefore, the physical attributes of biogenic habitats, here kelp holdfasts that better predict patterns in the assemblage structure of associated fauna can differ according to their different physical morphology, even though the overall assemblage structure of associated fauna was similar.     

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.     

Are kelp holdfasts islands on the ocean floor? – indication for temporarily closed aggregations of peracarid crustaceans

During the colonisation process of islands, newly immigrating species often arrive as single individuals. Islands that have received single colonisers may subsequently harbour large populations of a species, while other islands may completely lack this species. Exchange between islands is limited, thereby strongly affecting evolutionary processes. While this concept is widely used in the context of oceanic islands or habitat patches on the mainland, it is rarely used to explain and examine the distribution patterns of marine invertebrates. Benthic marine organisms inhabiting patches with island-like features may also be restricted in their movements between patches. Once established in a patch, it may be more favourable to remain there rather than moving to another patch. Juveniles of species with direct development may recruit to the island patch of their parents. Herein, we examined the peracarid fauna in patches that have island-like features, i.e. kelp holdfasts. The number of peracarid species within an individual holdfast increased with its size. Similarly, the number of individuals per holdfast increased with holdfast size. However, several peracarid species showed a strongly aggregated distribution pattern, being highly abundant in some holdfasts and almost completely absent in others. Our results suggest that these aggregations of conspecifics may be a consequence of the peracarid reproductive biology: fully developed juveniles emerge from the female's marsupium and recruit to the immediate vicinity of their mother, showing little or no tendency to emigrate towards other patches. At present, while it is not known how long peracarid aggregations within kelp holdfasts persist, our data suggest that some juveniles may remain with the natal holdfast and possibly reproduce therein. It is concluded that, during certain time periods, reproduction rates of peracarids in a holdfast may exceed their migration rates between holdfasts.     

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.     

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 Influence of Dry-season Conditions on the Bottom Dwelling Fauna of an East Australian Sub-tropical Estuary

Spatial patterns in benthic infaunal community structure of Port Curtis estuary (north-eastern Australia) were determined from quantitative grab samples and examined in relation to environmental variables. A total of 149 riverine, estuarine and open coastal stations were sampled during the winter survey, and 5744 individuals from 466 species identified. Filter-feeding organisms (primarily polychaetes, molluscs and crustaceans) dominated the benthos, and accounted for 50% of the total species abundance and 30% of the total species richness. Most taxa were uncommon, and 98% of species individually represented less than 2% of the total abundance. Distributional patterns in total species richness and abundance were highly correlated, and both parameters varied significantly with sediment grain size. Numbers of species and individuals were typically highest in coarse-sand and gravel sediments, and were lowest in fine, well-sorted, sands. Cluster analysis of species abundance data revealed nine community groupings characterised by small species sub-sets with restricted distributions. These groupings were primarily related to sediment grain size, sediment organic carbon content, salinity and depth. Not all variation in community structure was explained by these environmental variables, and it is likely that other unmeasured factors play an important role in determining benthic faunal composition in the estuary.     

Characterization of the Adhesive Holdfast of Marine and Freshwater Caulobacters

Caulobacters are prosthecate (stalked) bacteria that elaborate an attachment organelle called a holdfast at the tip of the cellular stalk. We examined the binding of lectins to the holdfasts of 16 marine Caulobacter strains and 10 freshwater species or strains by using a panel of fluorescein-conjugated lectins and fluorescence microscopy. The holdfasts of all the marine isolates bound to only wheat germ agglutinin (WGA) and other lectins that bind N-acetylglucosamine (GlcNac) residues. The freshwater caulobacters showed more variability in holdfast composition. Some bound only to WGA and comparable lectins as the marine strains did. Others bound additional or other lectins, and some did not bind to the lectins tested. The binding of WGA appeared to involve the regions of the holdfast involved with adhesion; a holdfast bound to WGA was significantly less adhesive to glass. Competition experiments with WGA-binding holdfasts and oligomers of GlcNac demonstrated that trimers of GlcNac (the preferred substrate for WGA binding) were more effective than dimers or monomers in preventing WGA binding to holdfasts, suggesting that stretches of contiguous GlcNac residues occur in the WGA-binding holdfasts. In addition, differences between freshwater and marine holdfasts in the strength of WGA binding were noted. The effect of a number of proteolytic and glycolytic enzymes on holdfast integrity was examined; the proteases had no effect for all caulobacters. None of the glycolytic enzymes had an effect on marine caulobacter holdfasts, but chitinase and lysozyme (both attack oligomers of GlcNac) disrupted the holdfasts of those freshwater caulobacters that bound WGA. Despite some similarity to chitin, holdfasts did not bind Calcofluor and no measurable effects on holdfast production were detectable after cell growth in the presence of diflubenzuron or polyoxin D, inhibitors of chitin synthesis in other systems. Finally, the holdfasts of all caulobacters bound to colloidal gold particles, without regard to the coating used to stabilize the gold particles. This binding was stronger or more specific than WGA binding; treatment with colloidal gold particles prevented WGA binding, but the reverse was not the case.     

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.     

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.     

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.     

Ediacaran macroalgal holdfasts and their evolution: a case study from China

A macroalgal holdfast (root-like structure) anchored or grown into sediments is a key trait of metaphytes and eukaryotic algae. Various patterns and taphonomic variants of congeneric holdfasts are preserved on the bedding planes of black shales of the Ediacaran Wenghui biota in South China. The macroalgal holdfast, which commonly consists of a rhizome, rhizoid and pith (perhaps mechanical tissue), can be morphologically classified into ten types within four groups of rhizomes: bare rhizome holdfasts (Grypania, Tongrenphyton and Sectoralga-type holdfasts), canopy rhizome holdfasts (Gemmaphyton, Gesinella, Discusphyton and Baculiphyca-type holdfasts), pithy rhizome holdfasts (Zhongbaodaophyton and pithy-cone-type holdfasts) and differentiated rhizome holdfasts (Wenghuiphyton-type holdfasts). Analysis of the Precambrian macroalgal record indicates that rhizomes played a more important role in the evolution of macroalgal holdfast than rhizoids. The following evolutionary stages of development of macroalgal holdfasts are proposed: (1) growth of the basal thallus into sediments (Grypania-type holdfast); (2) development of a primitive (indistinct) rhizome from the base of a stipe or thallus (Tongrenphyton-type and Sectoralga-type holdfasts); (3) growth of a distinct rhizome with rhizoid (Gemmaphyton-type, Gesinella-type and Discusphyton-type holdfasts); (4) development of a pith in the stipe (Baculiphyca-type holdfast); (5) pith extension into the rhizome (Zhongbaodaophyton-type and pithy-cone-type holdfasts); and (6) rhizome differentiation and development of a complex holdfast system (Wenghuiphyton-type holdfast).     

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.     

Community structure of vestimentiferan-generated habitat islands from Gulf of Mexico cold seeps

Biologically generated structures create habitat and influence the distribution and abundance of species in many marine systems. In the rather monotonous and nutrient-poor environment of the deep-sea, cold seep environments and their associated chemosynthetic communities offer islands of primary production and habitat to a generally sparsely distributed macrofauna. In this study, we investigate the structure of macrofaunal assemblages associated with vestimentiferan aggregations on the upper Louisiana slope of the Gulf of Mexico and the relationships between assemblage composition and the size and complexity of the vestimentiferan-generated habitat. Using custom-designed and custom-built devices, we collected seven whole vestimentiferan aggregations along with their associated fauna during the summers of 1997 and 1998. Sixty-five species were found associated with the four vestimentiferan aggregations collected in 1998, more than doubling the number of species previously reported for seeps in this region. Individual aggregations contained between 23 and 44 different non-vestimentiferan species. General trends of increasing species richness with increasing habitat size and increasing faunal density with increasing habitat complexity were identified, but substantial variability suggested other factors also control the composition of faunal associates. Faunal abundances decreased with increasing aggregation age. Seep endemics dominated the communities of younger aggregations, but non-endemic species dominated communities of older aggregations. Relative dominance of the heterotrophic community by primary consumers decreased, while predatory secondary and higher-order consumers increased with increasing aggregation age. These trends are discussed in terms of successional changes in aggregation structure, habitat heterogeneity and environmental conditions.     

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.     

Artificial substrata in a shallow sublittoral habitat: do they adequately represent natural habitats or the local species pool?

Artificial substrata have been advocated as tools which have considerable potential for monitoring both natural and anthropogenic effects on invertebrate communities of shallow coastal environments. In this experiment, community structure was compared between two dominant natural algal habitats (kelp holdfasts and algal turf) and artificial substratum units (ASUs; nests of pan scourers) deployed in close contact with, and 20 cm above the substratum. Univariate and multivariate statistical analyses were applied to the data to determine the similarity of community structure between the four different habitats. In addition, recently developed measures of taxonomic distinctness were applied to the data from both sets of artificial substrata to determine if they provided a representative sample of the local epifaunal species pool and thus have the potential to be used as surrogate samples for this important faunal group. There were marked differences between community structure in each of the habitats. Both sets of artificial substrata were dominated by tubicolous polychaetes with abundances that were more than an order of magnitude greater than in the holdfast and turf samples. The fauna recruiting to the artificial substrata deployed above the substratum showed the lowest values in the univariate summaries of diversity and evenness and were unrepresentative of the local species pool. Artificial samples deployed in contact with the substratum showed greater diversity and evenness but were still mostly unrepresentative of the local species pool. The tendency for both sets of artificial substrata to under-sample amphipods and to be dominated by suspension-feeding polychaetes suggests that methods using these units may be relatively insensitive to the effects of anthropogenic impacts (e.g. sewage outfalls) where shifts in community structure including increased dominance of suspension-feeders and polychaetes and a reduced dominance of amphipods have been observed. Further studies, including the evaluation of temporal variation in community structure related to the time at which the ASUs are deployed and duration of deployment, are needed to test the wider utility of artificial substrata as tools for monitoring shallow, sublittoral, epifaunal communities.     

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.     

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.     

Differences in response between haploid and diploid isomorphic phases of Gracilaria verrucosa (Rhodophyta: Gigartinales) exposed to artificial environmental conditions

This study tests the responses of juvenile gametophytes and tetrasporophytes (holdfast stage) of the isomorphic alga Gracilaria verrucosa under different environmental conditions.Estimations of survival and growth of holdfasts of haploid and diploid juvenile individuals were performed in natural sea-water and artificial culture medium, and under stringent conditions using lead as a toxin and ultra violet radiation as a mutagen. Results indicate that (i) holdfasts of haploid juveniles grow better than diploids in non-optimal medium conditions; (ii) holdfasts of diploid juveniles have a better tolerance to lead than haploids; and (iii) slight advantage of holdfasts of diploid juveniles grow better than haploids under U.V. radiation.     

Variations in holdfast attachment mechanics with developmental stage, substratum-type, season, and wave-exposure for the intertidal kelp species Hedophyllum sessile (C. Agardh) Setchell

Biomechanical models that describe physical and biological interactions on wave-exposed shores typically assume that a species' attachment properties are similar between seasons and sites. We tested this assumption using Hedophyllum sessile to investigate how macroalgal biomechanical attachment properties vary with developmental stage, substratum-type, season, and wave-exposure. Hedophyllum sessile is an intertidal kelp species that is able to survive in wave-exposed areas in the Northeast Pacific. For both juveniles and adults, holdfast attachment force and strength were measured at a wave-exposed and wave-protected site in Barkley Sound, British Columbia, Canada. Substratum and wave-exposure effects on attachment properties were tested in juvenile populations. Adult populations were sampled prior to (in July 1996) and after (in November 1996) a series of storms. Site and seasonal wave-exposure effects on attachment properties were tested in these adult populations. Comparisons to known attachment properties of other temperate macroalgal species were also made. Causes for these patterns are discussed but were not isolated in these studies. Juveniles' attachment properties differed on different substrata types and between wave-exposures, with the highest attachment forces and the most attached juveniles in articulated coralline algal turfs. Adult attachment is firm ( approximately 100 N), but relatively weak ( approximately 0.07 MNm(-2)). Adult attachment did not vary with site wave-exposure, but there was a shift within each site to more resistant holdfasts after a series of early winter storms. Seasonal increases in storm swells correlated to more thallus tattering and selected against large, loose holdfasts. The data presented here suggest that results from holdfast attachment field studies in one season cannot be extrapolated to another due to a complex set of dynamics. This is the first documentation of seasonal patterns in macroalgal attachment properties.