Spatially variable synergistic effects of disturbance and additional nutrients on kelp recruitment and recovery

Understanding the impact of multiple stressors on ecosystems is of pronounced importance, particularly when one or more of those stressors is anthropogenic. Here we investigated the role of physical disturbance and increased nutrients on reefs dominated by the canopy-forming kelp Ecklonia radiata. We combined experimental kelp canopy removals and additional nutrient at three different locations in a large embayment in temperate southeastern Australia. Over the following winter recruitment season, Ecklonia recruitment was unaffected by increased nutrients alone, but tripled at all sites where the canopy had been removed. At one site, the combination of disturbance and increased nutrients resulted in more than four times the recruitment of the introduced kelp Undaria pinnatifida. Six months after disturbance, the proliferation of the Undaria canopy in the canopy-removal and nutrient-addition treatment negatively influenced the recovery of the native kelp Ecklonia. Given the otherwise competitive dominance of adult Ecklonia, this provides a mechanism whereby Undaria could maintain open space for the following recruitment season. This interplay between disturbance, nutrients and the response of native and invasive species makes a compelling case for how a combination of factors can influence species dynamics.     

A molecular investigation of the genus Ecklonia (Phaeophyceae,Laminariales) with special focus on the Southern Hemisphere

Brown algae of the order Laminariales, commonly referred to as kelps, are the largest and most productive primary producers in the coastal inshore environment. The genus Ecklonia (Lessoniaceae, Phaeophyceae) consists of seven species with four species in the Northern Hemisphere and three in the Southern Hemisphere. It was recently transferred to the family Lessoniaceae based on phylogenetic analyses of nuclear and chloroplastic markers, though the type of the genus was not included and its relationship with allied genera Eckloniopsis and Eisenia remained unresolved. The present study is the first to produce a phylogeny focussed on the genus Ecklonia. It included sequences from nuclear, mitochondrial, and chloroplastic DNA, for most of the distribution range of the three current Southern Hemisphere species (Ecklonia radiata, Ecklonia maxima, and a sample of a putative Ecklonia brevipes specimen), sequences for East Asiatic species (Ecklonia cava, Ecklonia kurome, and Ecklonia stolonifera), as well as the closely related genera Eckloniopsis and Eisenia. Results confirmed E. radiata and E. maxima as two distinct species in South Africa, E. radiata as a single species throughout the Southern Hemisphere (in South Africa, Australia, and New Zealand) and East Asiatic species as a distinct lineage from the Southern Hemisphere clade. Results further pointed out a close sister relationship between Eckloniopsis radicosa and two Eisenia species (including the type species: Eisenia arborea) to the genus Ecklonia suggesting that the genera Eckloniopsis and Eisenia are superfluous.     

Experimental effects of kelp canopies on subtidal coralline algae

Plants are often grouped as canopy species or understorey species because it is thought that that these sets of taxa interact in predictable ways. Mensurative experiments in southern Australia demonstrated that the percentage cover of encrusting coralline algae was greater, and articulated (branching) coralline algae less, on boulders under a canopy of dense kelp (>7 plants per m²), Ecklonia radiata, than on boulders without kelp. Experimental clearances of kelp and reciprocal transplants of boulders between patches of E. radiata and patches without kelp showed that canopies maintained and facilitated the growth of encrusting coralline algae and reduced the cover of articulated coralline algae. Potential artefacts associated with clearing kelp and transplanting boulders were not detected when tested with a series of translocation controls. These results reject the model that the co‐occurrence of E. radiata and encrusting corallines is just an assemblage of plants caused by spatial and temporal coincidence. Instead, they support the model that kelp facilitates the growth and survival of understorey algae.     

Variation among diets in discrimination of δC and δN in the amphipod Allorchestes compressa

Discrimination of stable isotopes of carbon (δ¹³C) and nitrogen (δ¹⁵N) was examined for the amphipod Allorchestes compressa Dana using controlled laboratory experiments. Amphipods were fed exclusively on single diets (fresh or decomposed macroalgae or seagrass) for three weeks. Macrophyte type (i.e. seagrass, brown algae or red algae) had a greater influence on the stable isotope ratios of A. compressa than the state of decomposition of the macrophyte material. The experiments revealed that δ¹³C in A. compressa stabilised at values lower than those of the diets, which contrasts to the general assumption that consumer-diet discrimination of δ¹³C ranges from 0 to + 1‰. Amphipods fed on seagrass yielded the lowest δ¹³C values, which were 9 to 10‰ lower than their diet, while amphipods fed on macroalgae had values 2 to 4‰ lower than their diet. In addition, contrary to the general assumption that consumer-diet discrimination of δ¹⁵N ranges from + 3 to + 5‰, discrimination of δ¹⁵N was as low as − 1 and + 1 when A. compressa was fed on brown and red algae, respectively, but as high as + 3‰ when fed on seagrass. The results show that discrimination of stable isotopes of carbon and nitrogen can vary considerably depending on the food source, demonstrating that validation of assumptions about discrimination are critical for interpreting stable isotope data from field studies.     

Ecosystem engineering by a canopy‐forming kelp facilitates the recruitment of native oysters

Ecosystem engineers are species that influence the abiotic and biotic environment around them and may assist the restoration of associated species, including other habitat‐forming species. We deployed an array of 28 artificial reefs with transplanted Ecklonia radiata, the dominant canopy‐forming kelp species across southern Australia, to investigate how the patch size and density of E. radiata influenced the establishment of the associated communities of plants and animals. Many of the reefs were rapidly colonized by Ostrea angasi, a critically depleted reef‐forming oyster. Over the 24‐month deployment of the reefs, thick oyster mats formed across the entire surface of many of the reefs with estimated biomass densities exceeding 5 kg of live oysters/m²; however, oyster density was dependent on E. radiata patch size and density. Increasing patch size and the presence of kelp resulted in significantly higher densities of oysters 5 months after the reefs were deployed and at the end of the experiment, where oysters were approximately three times more numerous on reefs with kelp compared to those without kelp. E. radiata appeared to facilitate the establishment of O. angasi largely through its capacity to reduce benthic light and thus suppress competition from turfing algae. These results may inform the development of novel approaches to tackle recruitment bottlenecks affecting the restoration of O. angasi reefs.     

Survival of kelp adjacent to areas grazed by sea urchins in New South Wales,Australia

Abstract The sea urchin Centrostephanus rodgersii grazes areas of reef free of large brown algae (the barrens habitat). Survival of the kelp Ecklonia radiata adjacent to patches of barrens habitat was compared to that in the centre of kelp forests (centre) and edges of forests not adjacent to patches of barrens habitat (ungrazed). Estimates of rates of instantaneous mortality for tagged plants, as described by the slope of a negative exponential model (± 95% CI) were: barrens, 0.078 ± 0.004; centre, 0.051 ± 0.004 and ungrazed edge, 0.065 ± 0.007. Survival of plants was greatest in the middle of forests and least on the margins of patches of barrens habitat. A significant proportion of mortality in the barrens and centre positions was caused by herbivorous fish. When these plants were excluded from analysis there were no significant differences in survival between the barrens and ungrazed positions. It is concluded that C. rodgersii has little impact on the abundance of Ecklonia outside sharply defined boundaries.     

Characterisation of microsatellite loci in the habitat-forming kelp, Ecklonia radiata (Phaeophyceae, Laminariales)

Habitat forming algae play an important role in the ecology of temperate reefs worldwide. Despite this, our understanding of levels of gene flow within and among populations of algae is largely limited to studies on intertidal species; we know comparatively little about important habitat-forming subtidal algae. Here, we develop eight polymorphic microsatellite markers for the characterisation of population genetic diversity and structure in the subtidal kelp, Ecklonia radiata. This large macroalga is the most abundant habitat-forming kelp on the subtidal rocky reefs of temperate Australia and New Zealand where it forms extensive forests that support an astounding diversity of associated taxa.     

Modification of the physical environment by an <Emphasis Type="Italic">Ecklonia radiata</Emphasis> (Laminariales) canopy and implications for associated foliose algae

Macroalgal canopies modify their surrounding environment and thereby influence the structure of associated algal assemblages. Canopies can modify many factors that can be hard to separate and, consequently, the importance of individual factors often remains unknown. Experiments were carried out to test the hypotheses that Ecklonia radiata canopies modify light, sediment cover and water motion, and that each of these physical factors separately influence the assemblage of associated foliose algae. We measured light, sediment cover and water motion across six naturally occurring E. radiata densities and found a reduction in light and sediment cover as kelp density increased. The outcome for water motion was inconclusive. We also manipulated each of these three factors, while controlling for the two others, to determine the separate effects of light, sediment cover and water motion on the assemblage of foliose algae. Reduction in light had a strong effect on the foliose assemblage, reducing species richness and biomass. Reduction in sediment cover and water motion did not cause separate effects at the level of the assemblage, but the biomass of individual species of foliose algae indicated both positive and negative effects. We conclude that E. radiata canopies at Marmion, Western Australia, modify at least two factors of their physical environment, light and sediment cover. However, only light is modified to an extent where it has effects at the assemblage-level because, in contrast to the effects of sediment cover and water motion, the direction of responses are consistent among individual species of algae.     

Disturbance and reef topography maintain high local diversity in Ecklonia radiata kelp forests

Disturbance of competitive‐dominant plant and algae canopies often lead to increased diversity of the assemblage. Kelp forests, particularly those of temperate Western Australia, are habitats with high alpha diversity. This study investigated the roles of broad‐scale canopy loss and local scale reef topography on structuring the kelp‐dominated macroalgal forests in Western Australia. Eighteen 314 m² circular areas were cleared of their Ecklonia radiata canopy and eighteen controls were established across three locations. The patterns of macroalgal recolonisation in replicate clearances were observed over a 34 month period. Macroalgal species richness initially increased after canopy removal with a turf of filamentous and foliose macroalgae dominating cleared areas for up to seven months. A dense Sargassum canopy dominated cleared areas from 11 to 22 months. By 34 months, partial recovery of the kelp canopy into cleared areas had occurred. Some cleared areas did not follow this trajectory but remained dominated by turfing, foliose and filamentous algae. As kelp canopies developed, the initial high species diversity declined but still remained elevated relative to undisturbed controls, even after 34 months. More complex reef topography was associated with greater variability in the algal assemblage between replicate quadrats suggesting colonising algae had a greater choice of microhabitats available to them on topographically complex reefs. Shading by canopies of either Sargassum spp. and E. radiata are proposed to highly influence the abundance of algae through competitive exclusion that is relaxed by disturbance of the canopy. Disturbance of the canopy in E. radiata kelp forests created a mosaic of different patch types (turf, Sargassum‐dominated, kelp‐dominated). These patch types were both transient and stable over the 34 months of this study, and are a potential contemporary process that maintains high species diversity in temperate kelp‐dominated reefs.     

Patch formation by herbivorous fish in a temperate Australian kelp forest

Summary The odacid fish Odax cyanomelas feeds on the kelp Ecklonia radiata, an important component of subtidal reef habitats on the central coast of New South Wales, Australia. Herbivory by Odax has a major impact on the structure and dynamics of discrete patches within larger stands of kelp at Cape Banks. This three-year study showed that each year, between August and October, approximately the same patches of kelp were denuded by preferential feeding on the meristem and primary laminae. This coincided with a variable pulse of Ecklonia recruits to the cleared patches, thereby generating patches of a single age-class of plants. Neighbouring areas of Ecklonia forest, non cleared by Odax, consisted of larger, perennial plants, which exhibited more gradual changes in abundance. The seasonal impact of Odax appeared to be due to a change in the behaviour of female Odax during their spawning period. Observations suggested that females aggregate at traditional sites prior to spawning with territorial males in adjacent areas of kelp forest. An alternative hypothesis, that Odax preferentially attacked stands of one-year old Ecklonia plants, was rejected by a field experiment; the establishment of experimental stands of one-year old plants did not lead to increased damage due to Odax or any change in the use of space by the fish. The generality of this effect of fish herbivory is unknown, but this and other Odax species are widely distributed throughout temperate Australia, where Ecklonia is the dominant laminarian alga. The effects of pulsed herbivory by Odax is contrasted to the more continuous grazing by sea urchins in the same system. The latter herbivore has been shown to maintain areas free of Ecklonia, the long-term effects of herbivory by Odax remain unclear.     

Seasonal variation in the relationship between growth rate and phlorotannin production in the kelp Ecklonia radiata

Most theories for the evolution of plant chemical defences assume that defences are costly. In order to investigate the potential cost of phlorotannin production in the brown alga (kelp) Ecklonia radiata, I measured the correlation between changes in phlorotannin levels and growth rates for E. radiata in two seasons, spring and autumn. In spring, when both phlorotannin levels and growth rates in E. radiata were highest, there was a significant negative relationship between changes in phlorotannin levels and growth, consistent with a cost of phlorotannin production. No relationship was evident in autumn, indicating that moderate levels of phlorotannins and growth could be maintained simultaneously. These and related results suggest that the cost of defence will vary as a function of (1) endogenous patterns of growth or metabolite production and (2) variation in available resources. Evidence for a cost of phlorotannins in E. radiata adds to the paradox of the maintenance of high levels of phlorotannins in Australasian brown algae.     

Differentiation of benthic marine primary producers using stable isotopes and fatty acids: Implications to food web studies

A two-dimensional biomarker approach, using stable isotopes (δ¹³C, δ¹⁵N) and fatty acids, was used to evaluate differences both amongst and within benthic primary producer types (seagrass, fleshy red algae, calcareous red algae, brown algae, and seagrass periphyton) that are typical of the nearshore, temperate Australian region. The primary source of variance (as examined by permutational ANOVA) for all biomarkers examined was amongst primary producer types, as opposed to amongst species within type. δ¹³C showed a clear separation (Monte Carlo p < 0.05) between seagrass (range of means = −10.1 to −14.0‰) and macroalgae (−14.6 to −25.2‰), but could not differentiate amongst the algal types examined. Similarly, distinct δ¹⁵N signatures (p < 0.05) were found only for seagrass (range of means = 3.6-4.1‰) versus calcareous red algae (4.6-5.5‰), with all other types overlapping in their mean δ¹⁵N values. In contrast, multivariate analysis of fatty acid data (using Canonical Analysis of Principal coordinates; CAP) distinguished not only between seagrass and macroalgae, but also between red and brown algae (and to a limited extent between the calcareous and fleshy red algal types). The principal unsaturated fatty acids in the samples were C₂₀ polyunsaturates (found primarily in the macroalgae and periphyton), and C₁₈ mono- and polyunsaturates, with high proportions of 18:2n-6 and 18:3n-3 typical of the seagrasses. The C₁₈ monounsaturate 18:1n-7 was one of the most diagnostic compounds for the red algae examined, being present in very low amounts in seagrass and virtually absent in the brown algae. Conversely, brown algae were high in 18:4n-3, with 20:4n-3 particularly diagnostic of the kelp Ecklonia radiata. In contrast to stable isotopes, fatty acids helped distinguish different algal groups, thereby providing support that a two-dimensional approach using stable isotopes and fatty acids is likely to provide the most useful tool to distinguish primary producers in food web structure.     

Standing stock and production of Ecklonia radiata (C.Ag.): J. Agardh

The monthly productivity, standing stock, plant size and density of Ecklonia radiata (C.Ag.) J. Agardh is presented for a 2-yr period. Annual production was 20.7 kg wet wt · m^?2 with maximum growth of 0.9% per day in spring (October–December) and minimum growth of 0.2% per day in late summer. (March–April). A close negative correlation was found between spring and summer growth and water temperature. Maximum biomass (18 kg wet wt · m ^?2) did not coincide with maximum growth but occurred in late summer. Minimum biomass (6 kg wet wt · m ^?2) occurred in winter. An estimate of erosion of plant material from the kelp bed was made from these data and a hypothesis concerning the ultimate destination of eroded and removed kelp plants was formulated.     

CONTRIBUTION OF TEMPORAL AND SPATIAL COMPONENTS TO MORPHOLOGICAL VARIATION IN THE KELP ECKLONIA (LAMINARIALES)1

Environmental conditions that are known to cause morphological variation in algae (e.g., wave exposure) often vary in both space and time and are superimposed onto the distinct seasonal growth cycles of most temperate macroalgae. We tested the hypothesis that the morphology of the small kelp Ecklonia radiata (C. Agardh) J. Agardh is the product of an interaction between site (five reefs of different wave exposure) and the time of year that sampling occurs (summer vs. winter 2004). We determined that wave exposure had a strong directional effect on kelp morphology, with “Reefs” accounting for up to 43.4% of variation in individual morphological characters. “Times” had a narrowly nonsignificant effect on overall morphology but accounted for up to 31% of variation in individual characters. Many characters were affected by wave exposure, whereas only a few were (strongly) affected by time (e.g., thallus biomass). Interactive effects between “Reefs” and “Times” were generally small, accounting for 15.8% of variation in lamina thickness, but much less for most other characters. We conclude that wave exposure exerts a strong control over the morphology of E. radiata, but that the nature of the effect depends on the magnitude of wave exposure. We also conclude that most of the effects of wave exposure are consistent through time and do not interact with cycles of growth and pruning in any major way.     

Cross-habitat impacts of species decline: response of estuarine sediment communities to changing detrital resources

Food webs of many ecosystems are sustained by organic matter from other habitats. Human activities and climatic change are increasingly modifying the quality and supply of these resources, yet for most ecosystems it is unknown how the taxonomic composition of organic matter influences community composition. Along the coastline of Sydney, Australia, the once abundant habitat-forming macroalga, Phyllospora comosa, is now locally extinct. Shallow reefs are now primarily occupied by Sargassum sp. and, to a lesser extent, the kelp Ecklonia radiata. We experimentally manipulated the supply of P. comosa, Sargassum sp. and E. radiata to estuarine sediments to assess responses by macroinvertebrate communities to: (1) changing the identity of the dominant detrital resource; and (2) varying the ratio of input of different macrophytes. Estuarine sediments dosed with P. comosa supported greater abundances of macroinvertebrates than sediments receiving Sargassum sp. or the kelp E. radiata. Whereas plots receiving Sargassum sp. or E. radiata had fewer macroinvertebrates than controls, plots receiving a moderate (120 g dry weight per m²) loading of P. comosa had more. Mixtures of detritus dominated by P. comosa supported similar macroinvertebrate communities to monocultures of the alga. Communities in sediments receiving detritus comprised of less than one-third P. comosa were, however, distinctly different. Our study provides evidence that the ecological ramifications of species decline can extend to spatially removed ecosystems, subsidised by allochthonous materials. Even prior to extinction of detrital sources, small changes in their provision of organic matter may alter the structure of subsidised communities.     

Survival of juvenile Ecklonia radiata sporophytes after canopy loss

The understorey beneath a canopy of the kelp Ecklonia radiata often contains juvenile sporophytes of the same species. When canopy disturbance occurs, these juvenile sporophytes are exposed to new environmental conditions. If these juvenile sporophytes survive these new conditions, then they become a ready source of kelps to rapidly form a new canopy. This study investigated the potential of pre-existing juvenile sporophytes of E. radiata to survive post-disturbance conditions and contribute to the rapid formation of a new canopy. The potential of canopy recovery by recruitment of kelp from zoospores was also investigated. These processes were studied at different times during the summer and on reefs ranging in topographic complexity from simple, flat reefs to highly complex, rugose reefs. By tagging juvenile sporophytes after the adult kelp canopy was removed and monitoring them through time, it was demonstrated that most juveniles (> 50%) survived the change in conditions after canopy loss, with some juveniles going on to become members of a new canopy. Approximately 6-47% of tagged sporophytes died within 3-4 days after canopy removal possibly due to excessive photoinhibition and photostress as demonstrated by changes in photosynthetic performance (decreased alpha values) of juveniles. The potential contribution of juvenile sporophytes to the rapid formation of a new canopy appears to be dependent on the timing of canopy removal with late summer-autumn canopy loss favouring faster recovery. Topographically complex reefs had less short-term (7 days) survival of juvenile sporophytes than topographically simple reefs; however this difference was not carried through to the long-term (6 months) abundance of adult kelp in experimental clearings, which was greater on topographically complex reefs. Clearly, juvenile sporophytes in arrested development under existing canopies of the small kelp E. radiata are important for the rapid recovery of the kelp canopy once adults are lost through physical disturbance.     

Family‐level variation in early life‐cycle traits of kelp

Temperate kelp forests (Laminarians) are threatened by temperature stress due to ocean warming and photoinhibition due to increased light associated with canopy loss. However, the potential for evolutionary adaptation in kelp to rapid climate change is not well known. This study examined family‐level variation in physiological and photosynthetic traits in the early life‐cycle stages of the ecologically important Australasian kelp Ecklonia radiata and the response of E. radiata families to different temperature and light environments using a family × environment design. There was strong family‐level variation in traits relating to morphology (surface area measures, branch length, branch count) and photosynthetic performance (F_v/F_m) in both haploid (gametophyte) and diploid (sporophyte) stages of the life‐cycle. Additionally, the presence of family × environment interactions showed that offspring from different families respond differently to temperature and light in the branch length of male gametophytes and oogonia surface area of female gametophytes. Negative responses to high temperatures were stronger for females vs. males. Our findings suggest E. radiata may be able to respond adaptively to climate change but studies partitioning the narrow vs. broad sense components of heritable variation are needed to establish the evolutionary potential of E. radiata to adapt under climate change.     

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.     

VARIATIONS IN GROWTH,EROSION, PRODUCTIVITY,AND MORPHOLOGY OF ECKLONIA RADIATA (ALARIACEAE; LAMINARIALES) ALONG A FJORD IN SOUTHERN NEW ZEALAND1

Spatial and temporal patterns of growth, erosion, productivity, and morphology of the dominant habitat‐forming kelp Ecklonia radiata (C. Agardh) J. Agardh were studied bimonthly over 1.5 years in a southern New Zealand fjord characterized by strong gradients in light and wave exposure. Spatial differences in growth were observed with rates at two outer coast, high‐light, wave‐exposed sites reaching 0.42 and 0.45 cm · d^?1, respectively, compared to 0.27 cm · d^?1 at an inner, more homogeneous site. Sporophyte productivity was similar among sites, although population productivity was greater at the outer sites due to population density being 5‐fold greater than at the inner site. It was expected that the inner site would have no pronounced seasonal pattern in growth and productivity due to its homogeneity; however, all three sites displayed maximum rates in late winter/spring and minimal in autumn. Growth rates were 2‐fold greater during the first growth period than the following year. This discrepancy was not correlated to inorganic nitrogen (N) levels, which remained low year‐round (<4 μM), and is likely a result of an interaction between light and temperature, and the photosynthetic capability of E. radiata. Variable pigment content indicated photoacclimation at the inner site. Morphological differences were observed between sites, with E. radiata from the inner site having longer, wider, thinner blades and longer stipes. While E. radiata displayed spatial differences in growth, erosion, productivity, and morphology, populations displayed no temporal differences. These results highlight the need for greater understanding of the mechanisms influencing kelp growth and productivity in a unique marine environment.     

Predicting understorey structure from the presence and composition of canopies: an assembly rule for marine algae

Assembly rules provide a useful framework for predicting patterns of community assembly under defined environmental conditions. Habitat created by canopy-forming algae (such as kelps) provides a promising system for identifying assembly rules because canopies typically have a large and predictable influence on understorey communities. Across >1,000 km of subtidal South Australian coastline, we identified natural associations between assemblages of understorey algae and (1) monospecific canopies of Ecklonia radiata, (2) canopies comprised of E. radiata mixed with Fucales (Cystophora spp. and Sargassum spp.), and (3) gaps among canopies of algae. We were able to recreate these associations with experimental tests that quantified the assembly of understorey algae among these three habitat types. We propose the assembly rule that understorey communities on subtidal rocky coast in South Australia will be (1) monopolised by encrusting coralline algae beneath monospecific canopies of E. radiata, (2) comprised of encrusting corallines, encrusting non-corallines, and sparse covers of articulated corallines, beneath mixed E. radiata-Fucales canopies, and (3) comprised of extensive covers of articulated corallines and filamentous turfs, as well as sparse covers of foliose algae and juvenile canopy-formers, within gaps. Consistencies between natural patterns and experimental effects demonstrate how algal canopies can act as a filter to limit the subsets of species from the locally available pool that are able to assemble beneath them. Moreover, the subsets of species that assemble to subtidal rocky substrata in South Australia appear to be predictable, given knowledge of the presence and composition of canopies incorporating E. radiata.