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.     

Effects of kelp canopies on bleaching and photosynthetic activity of encrusting coralline algae

Canopy-forming algae often coexist with an understorey of encrusting coralline algae that bleach following the loss of canopies. We tested the hypothesis that canopy loss causes a reduction in photosynthetic activity of encrusting coralline algae concomitant with their bleaching. When canopies were experimentally removed, corallines bleached and their photosynthetic activity was rapidly reduced to half their activity observed under canopies. This result prompted us to test, and subsequently accept, the hypothesis that exposure of understorey corallines to enhanced light intensity per se (simulation of canopy loss) acts as a mechanism that causes bleaching and reduced photosynthetic activity. Despite bleaching, encrusting corallines maintained reduced levels of photosynthetic activity, and this may explain why, under certain conditions, bleached corallines can persist in the absence of canopy-forming algae. Nevertheless, our data provide evidence that the positive association between canopy-forming algae and encrusting coralline algae is maintained because of shade provided by the canopy.     

Variations in the configuration of algae in subtidal forests: Implications for invertebrate assemblages

Abstract The ecology of Australia's most extensive canopy‐forming alga, Ecklonia radiata, is often studied with little regard as to whether it occurs in monospecific stands or as part of a mixed assemblage of canopy‐forming algae. We tested the hypothesis that E. radiata does not primarily occur as monospecific stands, rather it occurs more often in stands of mixed algae. At a 1‐m² scale we recognized three main configurations within forests of algae (hereafter called stands): E. radiata that occurs as (i) monospecific stands; (ii) clumps (four or more individuals together) surrounded by species of Fucales; or (iii) individual plants (or clusters of fewer than three plants) interspersed among species of Fucales. All three types of stand occurred in similar proportions (percentage cover) across two regions of Australia's southern coastline (Western and South Australia). We also tested the hypothesis that these three types of stands (identified at 1 m²) contain different assemblages of invertebrates associated with the holdfast of E. radiata. Assemblages of invertebrates varied between monospecific and interspersed stands, but not between monospecific and clumped stands. These results suggest that variation in the configuration of subtidal algae (stands measured at a 1‐m² scale) has the potential to influence the composition and abundance of associated biota. We suggest that although studies in stands of monospecific E. radiata may provide useful information for the majority of forests containing E. radiata (monospecific and clumped stands made up 65% of forests sampled), caution must be used when extrapolating to stands of mixed, interspersed algae (>31% of forests sampled).     

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.     

3-D distribution of nongeniculate corallinales: a case study from a reef crest of South Sinai (Red Sea,Egypt)

An innovative technique for the estimation of species and growth-form abundance of coralline algae, including information on their vitality, was adopted on the reef crest of Ras Nosrani and Coral Bay, South Sinai. Data of coralline abundance from visual census and collection of voucher specimens were plotted on a 3-D sketched representation of the horizontal and vertical planes of the reef crest and of its crannies. Coralline dominance at the two investigated sites was not significantly different, with values ranging from 8.55 and 10.06% on the vertical plane and from 5.3 to 7.17% on the horizontal plane. About one-third of total corallines of the South Sinai reef crest was located in crannies, where the algae are completely overlooked by routine field surveys. Pink to violet, healthy corallines with encrusting growth-form, mainly belonging to Hydrolithon onkodes and Neogoniolithon, with subordinate fruticose Lithophyllum kotschyanum dominated the reef crest at both sites. The fruticose growth form, usually associated with L. kotschyanum, was more common in the horizontal than in the vertical plane. Purple, healthy, encrusting Sporolithon uncommonly occurred. Whitish, gray or green unhealthy or dead corallines were more common on the horizontal plane at both sites, possibly resulting from excessive solar radiation.     

A subtidal transect in Jervis Bay,New South Wales

Long-term variations in the intertidal algal flora of the entire rock platform at Plantation Point, Jervis Bay, are described by May (1981). The study now reported presents similar long-term changes in the flora of a subtidal region of the same headland, observed within the same period of time. This first detailed report of subtidal macroalgal communities in New South Wales describes a several-year study of the benthic communities along a transect in the upper sublittoral region of a rocky headland at Plantation Point, Jervis Bay. Eighty-nine species of algae were recorded, five of which were previously unrecorded for New South Wales. The area studied is dominated by the large brown algae Ecklonia radiata and Phyllospora comosa, large areas of which were cleared periodically by storms. Turf, shade and crustose coralline algal communities also were present. Storms, seasonal variation and longer term changes all affected the abundance and distribution of the algal species growing along the transect and hence the floristic composition of the area.     

Patterns in shallow subtidal marine assemblages along the coast of New South Wales

Shallow subtidal areas of rocky reefs in central and southern New South Wales may best be described as a mosaic of habitats, the distributions of which are seemingly related to depth, wave exposure and a number of biological processes, particularly herbivory. The Fringe habitat is generally found only in the most shallow waters. Forests of the laminarian alga Ecklonia radiata are often found at intermediate depths. In deeper, or more sheltered water, sponges, ascidians and red algae are more abundant and the abundances of sea urchins and other invertebrate grazers decline. Overlying this broad-brush pattern are patches of crustose coralline algae (the Barrens habitat), the distributions of which are not clearly related to depth. Invertebrate herbivores, and sea urchins in particular, are abundant in the Barrens habitat. The Barrens habitat was most represented at the more southern locations. At the two most northern locations, reefs were shorter in length and dominated by ascidians (Pyura species).     

Loss of consumers alters the effects of resident assemblages on the local spread of an introduced macroalga

Despite the great interest for the role played by resident assemblages in regulating biological invasions, few studies have assessed how these can influence the spread of exotic species that have successfully established or have included more than one trophic level. On shallow rocky reefs, we assessed how the effects of different benthic assemblages on the spread of an invasive alga, Caulerpa racemosa, are influenced by alterations in the density and species composition of the resident sea urchin assemblage. In order to simulate herbivore species loss scenarios, assemblages dominated by different morphological groups of algae (i.e. turfs or encrusting corallines) or experimentally cleared plots (i.e. bare rock) were exposed to grazing by different combinations of species (Arbacia lixula and Paracentrotus lividus) and densities (natural, ?50% and ?100% of natural densities) of urchins. Algal turfs and encrusting corallines generally facilitated C. racemosa. Manipulating urchins assemblages did not affect the cover and density of fronds of C. racemosa. In contrast, halving the density of P. lividus favoured the penetration of stolons of C. racemosa, consistently among algal assemblages. Other effects of urchins varied among algal assemblages, indicating interactions between trophic levels. In algal turfs, the total removal of urchins caused a decrease in the penetration of stolons, while it enhanced the length of fronds, indicating a shift in the growth form of this clonal plant and, ultimately, a depression of its spreading ability. In bare or encrusting corallines dominated surfaces, the removal of urchins had positive effects on the penetration of stolons and on the length of fronds of C. racemosa, irrespective of one or two species being manipulated and for the intensity of their removal. Our results show that tradeoffs between negative and positive effects of herbivores, varying according to relative densities of species and to the direction and strength of the effects of resident plant assemblages, can influence local rates of spread of C. racemosa. Thus, not only facilitation of exotics by natives can be key in enhancing the spread of exotic species, but it can occur between organisms at different trophic levels. Finally, our findings have important implications for the management of C. racemosa, in view of the progressive domination of shallow rocky reefs by algal turfs.     

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.     

The grazing effect of limpets on the macroalgal community of a rocky intertidal shore

Abstract Abundance of macro-algae in the mid-littoral zone on a Victorian intertidal rocky shore varied seasonally, algae being rare in Summer and common in Winter and Spring. Field experiments demonstrated that, of two species of grazing limpets (Cellana tramoserica and Siphonana diemenensis), only Siphonaria had a major effect on the abundance of foliose algae (e.g. Scytopsiphon lomeniarid) and neither species had a great effect on the encrusting algae (Ralfsia spp.). There was no evidence of competition for food between the two species of limpets, in contrast with results found for similar species in New South Wales.     

Canopy manipulations of exotic Bitter Willow (Salix elaeagnos) forest for indigenous seedling recruitment: A pilot study

The invasive exotic tree species Bitter Willow (Salix elaeagnos; Salicaceae) has colonised areas of rank exotic grassland and has been found to contain indigenous seed, dispersed by frugivorous birds into the monospecific stands. This small pilot study examined whether indigenous seedlings that have germinated in the understorey of exotic Bitter Willow stands could be stimulated to establish through the creation of small‐scale canopy gaps. In Bitter Willow forest, four single Bitter Willow trees were poisoned to create canopy gaps. Light transmission and seedling regeneration of tree and shrub species were assessed beneath both the four manipulated and three comparable intact Bitter Willow canopies. Over 3 years, seedling height and density increased more beneath opened compared to intact Bitter Willow canopies. These results suggest that Bitter Willow can fill the roles of both a facilitative nurse and a perch tree. Larger‐scale canopy manipulation experiments of both Bitter Willow and other Salix species are needed to determine the full potential of canopy manipulations for forest restoration.     

Recruitment of the intertidal kelp Lessonia nigrescens Bory in northern Chile: successional constraints and opportunities

The strong 1982/83 El Niño event caused local extinction in populations of the low intertidal kelp Lessonia nigrescens Bory on northern Chilean coasts. The kelp has partially recolonized, but its recovery has apparently been low. This study documents the effect of biological factors that potentially decrease the velocity of kelp recolonization in northern Chile. A removal experiment showed that encrusting coralline algae, which dominate the lower intertidal, significantly reduce the recruitment of L. nigrescens in the presence of herbivores. Epithallium shedding by encrusting corallines is the most probable cause for this inhibition process. On the other hand, grazing on encrusting corallines by the chiton Enoplochiton niger (Barnes), a large-sized herbivore (10–20 cm length), could also affect kelp recruitment. The intestinal content of E. niger, where encrusting corallines are the main item (84.2%), revealed the presence of L. nigrescens. Among the natural substrata on which kelp recruits, measured at five different localities, the coarsely-branched alga Corallina officinalis (Decaisne) Kützing reached the highest frequency, despite its extremely low cover (< 1%) in the field. This suggests that settlement on turfs of C. officinalis allows the kelp to escape from herbivory, thus facilitating its recruitment. The roles of timing of kelp recruitment and seasonal grazing are also discussed.     

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.     

Biodiversity and trophic structure of invertebrate assemblages associated with understorey red algae in a Laminaria digitata bed

Although the composition of invertebrate assemblages associated with kelps has motivated several studies in the recent past, little is known about assemblages associated with smaller, understorey macroalgae in these ecosystems. Here, the composition of invertebrate assemblages living within understorey macroalgae of a kelp (Laminaria digitata) forest has been studied over two sampling dates, and the structure of the food web investigated using stable isotopes. A total of 145 species of mobile fauna, mainly amphipods and gastropods, were identified. Although differences were date-dependent, we showed that the three species considered (Palmaria palmata, Mastocarpus stellatus, Ellisolandia elongata) sheltered different associated assemblages, including high host-specificity, which suggests that the animal diversity associated with rocky shores is enhanced by a high algal diversity. Overall, a dominance of gastropods was observed for the two-dimensional, leaf-like P. palmata, while the three-dimensionally structured species (M. stellatus, E. elongata) were characterized by a dominance of amphipods. Stable isotopes highlighted different trophic structures; E. elongata-associated assemblages were most likely relying on a dominant food source, sediment organic matter, while other assemblages were based on a wider diversity of food sources (algae, sediment, suspended organic matter). Our results illustrate the need to consider the microhabitats formed by understorey macroalgae in order to thoroughly assess the diversity and understand the functioning of coastal rocky ecosystems.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

Invasion of alien macroalgae in different Mediterranean habitats

The study evaluated different macroalgal invasions in the main Mediterranean coastal habitats on hard bottom. Biodiversity, species composition and structure of macroalgal assemblages were compared among non-invaded areas and areas invaded by the Chlorophyta Caulerpa racemosa var. cylindracea and by the turf-forming Rhodophyta Womersleyella setacea in three different habitats: shallow rocky bottom, deep rocky bottom and dead matte of the seagrass Posidonia oceanica. Results showed that alien macroalgae constituted a relevant component of benthic assemblages in invaded areas of the Mediterranean Sea. Assemblages invaded by Womersleyella setacea and Caulerpa racemosa showed lower values of diversity and large differences in the structure and species composition related to non-nvaded assemblages. The species that mostly suffered from invasion were erect species reproducing sexually; moreover, the dominance of W. setacea led to low abundance of native filamentous algae, while C. racemosa colonization seemed particularly threatening for encrusting algae. All the studied habitats appeared highly invasible by alien macroalgae, even if W. setacea appeared more invasive in deeper habitats, while colonization of C. racemosa seemed more serious in shallower habitats; the dead matte of P. oceanica represented a suitable substrate for the spread of both species. Differences among assemblages in different habitats were reduced in invaded areas.     

Kelp and sea urchin settlement mediated by biotic interactions with benthic coralline algal species

Species interactions can influence key ecological processes that support community assembly and composition. For example, coralline algae encompass extensive diversity and may play a major role in regime shifts from kelp forests to urchin-dominated barrens through their role in inducing invertebrate larval metamorphosis and influencing kelp spore settlement. In a series of laboratory experiments, we tested the hypothesis that different coralline communities facilitate the maintenance of either ecosystem state by either promoting or inhibiting early recruitment of kelps or urchins. Coralline algae significantly increased red urchin metamorphosis compared with a control, while they had varying effects on kelp settlement. Urchin metamorphosis and density of juvenile canopy kelps did not differ significantly across coralline species abundant in both kelp forests and urchin barrens, suggesting that recruitment of urchin and canopy kelps does not depend on specific corallines. Non-calcified fleshy red algal crusts promoted the highest mean urchin metamorphosis percentage and showed some of the lowest canopy kelp settlement. In contrast, settlement of one subcanopy kelp species was reduced on crustose corallines, but elevated on articulated corallines, suggesting that articulated corallines, typically absent in urchin barrens, may need to recover before this subcanopy kelp could return. Coralline species differed in surface bacterial microbiome composition; however, urchin metamorphosis was not significantly different when microbiomes were removed with antibiotics. Our results clarify the role played by coralline algal species in kelp forest community assembly and could have important implications for kelp forest recovery.     

Understorey vegetation moderates climate in open forests: The role of the skirt-forming grass tree Xanthorrhoea semiplana F.Muell

Microsites are created by abiotic and biotic features of the landscape and may provide essential habitats for the persistence of biota. Forest canopies and understorey plants may moderate wind and solar radiation to create microclimatic conditions that differ considerably from regional climates. Skirt-forming plants, where senescent leaves create hut-like cavities around the stem, create microsites that are sheltered from ambient conditions and extreme weather events, constituting potential refuges for wildlife. We investigate day and night temperatures and humidity for four locations (grass tree cavities, soil, 20 cm above-ground, 1 m above-ground) in a South Australian forest with relatively open canopy of stringybark eucalypts (Eucalyptus baxteri, E. obliqua) and an understorey of skirt-forming grass trees (Xanthorrhoea semiplana) at 5, 10, 20, and 40 m from the forest edge. We also measured the percentage of canopy and understorey covers. Generally, temperature and humidity differed significantly between more sheltered (grass tree cavities, soil) and open-air microsites, with the former being cooler during the day and warmer and more humid during the night. Furthermore, our results suggest that canopy cover tends to decrease, and understorey cover tends to increase, the temperature of microsites. Distance to the edge was not significantly related to temperature for any microsite, suggesting that the edge effect did not extend beyond 10 m from the edge. Overall, grass trees influenced microclimatic conditions by forming a dense understorey and providing cavities that are relatively insulated. The capacity of grass tree cavities to buffer external conditions increased linearly with ambient temperatures, by 0.46°C per degree increase in maximum and 0.25°C per degree decrease in minimum temperatures, potentially offsetting climate warming and enabling persistence of fauna within their thermal limits. These climate moderation properties will make grass trees increasingly important refuges as extreme weather events become more common under anthropogenic climate change.     

Relationship between algae and environment: an Early Cretaceous case study, Trascău Mountains, Romania

The relationship between algae and depositional environment was studied in a limestone succession of Berriasian–Valanginian age. Several depositional environments were recorded from shallow subtidal to intertidal and supratidal, with salinity ranging from normal marine to fresh and/or supersaline water. The algal assemblages consist mainly of dasycladaleans, rivulariacean-type cyanobacteria and charophytes. Nipponophycus (Bryopsydales) and Lithocodium-Bacinella (microproblematicum), occur at some levels. Dasycladaleans are associated with subtidal, sometimes restricted (“lagoonal”) environments, while rivulariacean-like cyanobacteria and charophytes characterise intertidal-supratidal and fresh and/or supersaline environments, respectively. Among the dasycladaleans, Salpingoporella annulata is often related to restricted environments where it forms monospecific assemblages. Large dasycladaleans, such as Selliporella neocomiensis, Macroporella praturloni and Pseudocymopolia jurassica are found in subtidal high-energy deposits (bioclastic grainstones). The relationship between environment and algae, characteristic for each depositional unit, can be used to interpret the relative sea-level variations.