Reproductive Output,Synchrony across Depth and Influence of Source Depth in the Development of Early Life stages of Kelp

Ecklonia radiata is the main foundation species in Australian temperate reefs, yet little has been published on its reproduction and how this may change across its depth range (1–50+ m). In this study, we examined differences in sporophyte morphology and zoospore production during a reproductive season and across four depths (7, 15, 25, and 40 m). Additionally, we examined differences in germination rate, survival, and morphological traits of gametophytes obtained from these four depths, cultured under the same light and temperature conditions. Multivariate morphology of sporophytes differed significantly between deep (~40 m) and shallow sites (7 and 15 m), but individual morphological traits were not significantly different across depths. Total spore production was similar across depths but the peak of zoospore release was observed in February at 15 m of depth (6,154 zoospores · mm^?2 of tissue) and the minimum observed in January at 7, 25, and 40 m (1,141, 987, and 214 zoospores · mm^?2 of tissue, respectively). The source depth of zoospores did not have an influence in the germination rate or the survival of gametophytes, and only gametophytes sourced from 40 m sites presented significantly less surface area and number of branches. Overall, these results indicate that E. radiata’s reproductive performance does not change across its depth range and that kelp beds reproducing in deeper areas may contribute to the replenishment of their shallow counterparts. We propose that deep kelps may constitute a mechanism of resilience against climate change and anthropogenic disturbances.     

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.     

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.     

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.     

Age- and size-based morphological comparison between the brown alga Sargassum macrocarpum (Heterokonta; Fucales) from different depths at an exposed coast in northern Kyoto, Japan

Although large perennial brown algae are known to show great morphological plasticity responding to specific environmental factors such as depth and wave exposure, there are few reports showing this morphological variability taking into account age or size composition. Here, we conducted age- and size-based morphological comparison between the perennial brown alga Sargassum macrocarpum C. Agardh from their upper depth limit, middle depth, and lower depth limit at an exposed coast in northern Kyoto, Japan. Model II regression was fitted for the relationships between age or stipe diameter (SD) and morphological variables including SD, holdfast weight (HW), number of main branches (MBN), total weight of main branches (MBW), thallus weight (TW), and thallus length (TL) of 30 specimens collected seasonally from each depth zone. The differences between depths in these regression equations were analyzed using SMATR. Although SD, HW, and MBN of the specimens did not differ between depths under both age- and SD-basis, there were significant differences between depths in MBW, TW, and TL, suggesting that the differences in TW between depths are resulted from differences in MBW. Whereas TW at the lower depth limit was lower than that at the upper depth limit or middle depth in many months, TL at the upper depth limit was shorter than that at the middle depth or lower depth limit in some months. These results suggest that S. macrocarpum at shallower depths tend to have short and bushy morphology, whereas those at deeper depths have long and less bushy morphology.     

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).     

External and internal factors regulating metabolic rates of an estuarine benthic community

Summary The structure and metabolism of a soft-sediment estuarine macrofaunal community were measured over an annual cycle at two depth-contours in mesohaline Chesapeake Bay. Additional data for plankton productivity and respiration, as well as seston and sediment organics are also summarized for these communities. Benthic community respiration ranged from 0.24–3.38 g O₂ m^-2 d^-1, and significant differences were detected between the two depths. Similarly, macroinfaunal standing stocks reached 11.2 and 32.3 g (ash free) m^-2 for 3 m and 6 m depth communities, respectively, and both exhibited mid-summer declines in abundance. Inferences drawn from these data facilitated a partitioning of benthic community respiration into macrofaunal and meiofaunal/microbial components with a residual term, much of which could be explained statistically by interactions between these two components. A multi-variate statistical model developed from these data matched benthic respiration measurements within 1–2 S.E. Mass-balances of organic carbon were estimated for water column and benthos at the two depthcontours for early and late summer, as well as for an entire, time-weighted year. These various analyses led to the tentative conclusions that this benthic community was regulated by such internal factors as macrofaunal/meiofaunal grazing and microbial gardening, and by external factors such as temperature and predation by nekton. However, it appears that the ultimate control for this community was the supply of energy from organic carbon.Contribution No. HPEL-1206, USASupported by grants with the Maryland Department of Natural Resources (PS-72-02(77-78)), J.A. Mihursky, Coordinating Principal Investigator     

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).     

Long-Term Observations of Epibenthic Fish Zonation in the Deep Northern Gulf of Mexico

A total of 172 bottom trawl/skimmer samples (183 to 3655-m depth) from three deep-sea studies, R/V Alaminos cruises (1964–1973), Northern Gulf of Mexico Continental Slope (NGoMCS) study (1983–1985) and Deep Gulf of Mexico Benthos (DGoMB) program (2000 to 2002), were compiled to examine temporal and large-scale changes in epibenthic fish species composition. Based on percent species shared among samples, faunal groups (≥10% species shared) consistently reoccurred over time on the shelf-break (ca. 200 m), upper-slope (ca. 300 to 500 m) and upper-to-mid slope (ca. 500 to 1500 m) depths. These similar depth groups also merged when the three studies were pooled together, suggesting that there has been no large-scale temporal change in depth zonation on the upper section of the continental margin. Permutational multivariate analysis of variance (PERMANOVA) also detected no significant species changes on the limited sites and areas that have been revisited across the studies (P>0.05). Based on the ordination of the species shared among samples, species replacement was a continuum along a depth or macrobenthos biomass gradient. Despite the well-known, close, negative relationship between water depth and macrofaunal biomass, the fish species changed more rapidly at depth shallower than 1,000 m, but the rate of change was surprisingly slow at the highest macrofaunal biomass (>100 mg C m⁻²), suggesting that the composition of epibenthic fishes was not altered in response to the extremely high macrofaunal biomass in the upper Mississippi and De Soto Submarine Canyons. An alternative is that the pattern of fish species turnover is related to the decline in macrofaunal biomass, the presumptive prey of the fish, along the depth gradient.     

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.     

盐城滨海滩涂湿地典型植物群落土壤微生物组成与结构特征

土壤微生物是生态系统维持正常结构与功能的重要组成部分,为探究盐城滩涂典型湿地土壤微生物群落结构特征,以江苏盐城滩涂互花米草、藨草、盐地碱蓬、芦苇及淤泥质光滩5种典型群落为对象,采用16S rRNA高通量测序技术分析0—10 cm(表层)、10—30 cm(中层)、30—60 cm(深层)土壤微生物多样性及群落结构。结果表明：(1)几种植物群落间,土壤微生物群落结构差异较大,主要体现在细菌群落结构的差异性,古菌群落结构差异相对较小。光滩与植物群落间,在土壤细菌种类及相对丰度上差异相对较大,互花米草群落与本土植物群落间,在微生物群落的细菌种类组成上存在较大差异;藨草群落土壤表层微生物群落结构与互花米草群落相似,深层与盐地碱蓬、芦苇群落相似。(2)同一群落不同层次土壤微生物群落结构相似,差异小于不同群落间土壤微生物群落的结构差异性;不同群落对应层次间,表深层土壤中五种群落土壤微生物多样性存在显著差异,中层土壤中五种群落微生物多样性差异不显著。总体上,植物群落类型对土壤微生物群落结构的影响大于土壤深度;与本土植物群落相比,互花米草群落土壤主要优势门微生物种类差异较小,但部分优势门微生物相对丰度...     

The effects of water depth and density on the growth of a unionid clam*

SUMMARY. 1. Unionid clams from Narrow Lake, Alberta, were collected to quantify the natural variation in growth, to assess the natural variation in abundance, age and size distribution, and growth with water depth in the lake, and to conduct in situ experiments to directly test the effects of water depth (temperature) and clam abundance on clam growth. 2. The unionid clam, Anodonta grandis simpsoniana, showed wide variation in length at a given age. There were no significant differences in growth between clams collected at 1,3, 5, and 7m depths in the lake despite marked differences in water temperature. The wide variation in clam biomass within each depth zone may have masked possible effects of water depth. 3. The effect of water depth and variation in clam density on clam growth was tested directly by stocking clams into small enclosures at densities equivalent to 50, 150, 250, 350 and 450g m-2 (live weight) at each of 1, 3, 5 and 7 m depths in Narrow Lake (each depth and abundance treatment in triplicate). A uniform sandy substrate was used in all enclosures to eliminate any possible effect of substrate type on growth. 4. Mortality was negligible (0.9%) during the experiment. Clam density had no significant effect on clam growth which suggests that clam growth was not food limited in the lake. 5. Clams reared at 7 m grew more slowly than clams reared at 1, 3 and 5 m. Clams reared at 5 m grew more slowly than clams reared at 1 and 3m. Growth of clams reared at 1 and 3m did not differ. These differences in growth were strongly correlated with the measured differences in water temperature between depths. 6. Migration between depths probably accounts for the lack of a depth effect on clams growing in the natural habitat.     

不同水深条件下刺参养殖池塘大型底栖动物群落结构季节性变化特征

于2012年7月至2013年4月调查了荣成靖海湾3个不同水深的刺参(Apostichopus japonicus)养殖池塘内大型底栖动物的构成,以了解不同水深对刺参养殖池塘环境条件的影响以及由此引起的大型底栖生物群落结构的改变。结果表明:3个不同水深梯度池塘(1#浅水位、2#正常水位和3#高水位)底部光照强度、叶绿素a(Chla)和总有机物(TOM)含量存在显著差异,各池塘水温差异不显著。光强、Chla和TOM含量在夏季、冬季和春季均表现为1#池塘显著高于3#池塘;秋季各池塘间光强和TOM含量差异不显著,Chla含量则表现为3#池塘显著高于1#池塘。各季节3个池塘间大型底栖动物在种类组成、丰度、生物量和多样性指数上均存在显著性差异。大型底栖动物丰度和生物量夏季均表现为1#池塘显著高于3#池塘,秋季和冬季则相反;春季1#池塘丰度显著高于3#池塘,生物量则差异不显著。这些差异主要与其各自优势种及其优势度指数大小有关。大型底栖动物多样性指数夏季和秋季均表现为1#池塘高于3#池塘,春季则相反,冬季各池塘间多样性指数差异不显著。单因子相似性分析(ANOSIM)表明,各季节3个池塘间大型底栖动物群落结构均存在显著差异,表明水深梯度对刺参养殖池塘大型底栖动物群落结构造成显著性影响。相似性百分比分析(SIMPER)显示,各季节对3个池塘间大型底栖动物群落差异起主要作用的物种为各个池塘的优势种。典范对应分析(CCA)表明,水深、Chla和TOM含量为影响大型底栖动物群落的主要环境因子。     

Rhodolith beds at the easternmost extreme of South America: Community structure of an endangered environment

Rhodolith beds are built by the aggregation of free living marine benthic coralline algae. Herein, we described phytobenthic communities associated with subtidal rhodolith beds in northeastern Brazil and tested the hypothesis that depth affects their structure. We compared macroalgal assemblages from depths of 10, 15 and 20 m. The genus Lithothamnion was dominant in these beds. Rhodolith density was similar at different depths, but volume decreases as depth increases. Sixty-seven species of fleshy algae were collected. The red algal order Ceramiales was dominant. A distinct community corresponds to each sampled depth. The shallower depth presented higher values for biomass, number of species, Shannon-Wiener diversity, and Pielou's evenness. When depth and water transparency increased, the number of species and the abundance of macroalgae decreased.     

Changes in the fish fauna associated with a sub-Antarctic <Emphasis Type="Italic">Macrocystis pyrifera</Emphasis> kelp forest in response to canopy removal

The fish fauna associated with a Macrocystis pyrifera forest and the effects of the canopy removal on this fauna were studied in the Beagle Channel, Tierra del Fuego. Seasonal changes and differences in the fish communities that inhabit the canopy and the holdfast were also studied. Two patches of kelp forest separated by 200 m were selected. In one, the canopy was removed periodically while the other was used as a control. Samples were collected seasonally, from autumn 1999 to 2001. Fish fauna in the water column was sampled using trammel nets and holdfast fish fauna was sampled by removing the complete holdfast. Different assemblages of fish species were captured in the water column (surface and bottom) and in the holdfast. In the former there were mainly pelagic and benthopelagic species and the latter predominantly demersal species. The principal effect of the canopy removal was a drop in the abundance (and total weight) of Paranotothenia magellanica (doradito), the species with the strongest relationship with the canopy, principally at the surface of the treated patch probably due to a reduction in the availability of refuges. A high degree of seasonality was observed for fish species’ total weight, abundance and diversity, with higher values in summer and autumn. Taking into account our data and available data on kelp growth in the same locality, we suggest of cutting the kelp forest (preferably those farther away from the coast) once a year in winter or early spring, in order to minimize impact on the fish community.     

Reef habitats and associated sessile-benthic and fish assemblages across a euphotic–mesophotic depth gradient in Isla Desecheo, Puerto Rico

Quantitative surveys of sessile benthos and fish populations associated with reef habitats across a 15–50 m depth gradient were performed by direct diver observations using rebreathers at Isla Desecheo, Puerto Rico. Statistically significant differences between depths were found for total live coral, total coral species, total benthic algae, total sponges and abiotic cover. Live coral cover was higher at the mid-shelf (20 m) and shelf-edge (25 m) stations, whereas benthic algae and sponges were the dominant sessile-benthic assemblage at mesophotic stations below 25 m. Marked shifts in the community structure of corals and benthic algae were observed across the depth gradient. A total of 119 diurnal, non-cryptic fish species were observed across the depth gradient, including 80 species distributed among 7,841 individuals counted within belt-transects. Fish species richness was positively correlated with live coral cover. However, the relationship between total fish abundance and live coral was weak. Abundance of several numerically dominant fish species varied independently from live coral cover and appeared to be more influenced by depth and/or habitat type. Statistically significant differences in the rank order of abundance of fish species at euphotic vs mesophotic stations were detected. A small assemblage of reef fishes that included the cherubfish, Centropyge argi, sunshine chromis, Chromis insolata, greenblotch parrotfish, Sparisoma atomarium, yellowcheek wrasse, Halichoeres cyanocephalus, sargassum triggerfish, Xanthichthys ringens, and the longsnout butterflyfish, Chaetodon aculeatus was most abundant or only present from stations deeper than 30 m, and thus appear to be indicator species of mesophotic habitats.     

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.     

Sublittoral Macrobenthic Infaunal Assemblages of Two Nearby Embayments from Central Chile

The benthic assemblages in two Central Chile nearby embayments were studied from quantitative samples collected from 15 sites at depths of 8–65 m. The macrobenthic infauna (<0.5 mm) of both bays was greatly dominated by polychaetes. Some 93 taxa were identified, of which 51 were polychaetes. The average macrofaunal abundance for all stations (15,021 ind. m⁻²) is very close to the values reported for the neighboring areas. Numerical classification and ordination (DCA) of sites resulted in three site-groups mostly reflecting differences in the bottom sediments: the muddy-bottom stations of Concepción Bay and the shelf-associated stations, the sandy-bottom stations of San Vicente Bay and an heavily polluted station at San Vicente port. Classification of species showed that the muddy-bottom stations and the sandy-bottom sites had characteristic species assemblages. The macrofaunal assemblages presented high dominance values, which were due to the high numerical abundances of a few species in the collections.     

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.