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Morphology of Ecklonia Radiata (Phaeophyta: Laminarales) Along Its Geographic Distribution in South-Western Australia and Australasia

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Morphology of Ecklonia Radiata (Phaeophyta: Laminarales) Along Its Geographic Distribution in South-Western Australia and Australasia Marine Biology (2003) 143: 47–55 DOI 10.1007/s00227-003-1069-9 T. Wernberg Æ M. Coleman Æ A. Fairhead Æ S. Miller M. Thomsen Morphology of Ecklonia radiata (Phaeophyta: Laminarales) along its geographic distribution in south-western Australia and Australasia Received: 18 November 2002 / Accepted: 7 January 2003 / Published online: 10 April 2003 Ó Springer-Verlag 2003 Abstract Ecklonia radiata (C. Ag.) J. Agardh is a com- thallus length was 135.2±12.5 cm in Kalbarri and only mon macroalga on reefs in the warm-temperate parts of 69.7±5.5 cm in Sydney. There were no consistent the southern hemisphere. It is a dominant habitat- spatial patterns of variation among individual morpho- former and as such has a strong structuring effect on logical characters, and, generally, variations among associated algal assemblages. Morphological variation individual characters were poorly correlated ()0.5 in E. radiata potentially affects its interactions with the <R<0.5). This suggests that individual morphological surroundings and contributes to confusion about its characters develop independently of each other in re- taxonomy. We quantified the magnitude of morpho- sponse to processes operating at different spatial scales. logical variation in fully developed E. radiata sporo- Multivariate measures of morphology were found to be phytes across Australasia and tested the hypotheses that different among some locations and similar among E. radiata has different morphology at different loca- others ()0.37 £ Clarke’s R £ 1), but there was no cor- tions and that the degree of morphological difference relation (Spearman’s R=0.08) between morphological depends on spatial distances among locations. A total of similarity and distance between locations. Consequently, 11 morphological characters were sampled from 11 lo- our results do not support clinal variation in E. radiata cations along the Australian coastline from Kalbarri in morphology. Rather, they suggest the presence of dis- Western Australia to Sydney in New South Wales as crete morphologically different populations, in which the well as from Doubtful Sound, New Zealand. Most morphology at any one location reflects multiple forcing morphological characters varied considerably from one factors operating on different morphological characters location to another. For example, the average (±SE) at different spatial scales. Communicated by G.F. Humphrey, Sydney Introduction T. Wernberg (&) School of Plant Biology, University of Western Australia, Ecklonia radiata (C. Ag.) J. Agardh is a small kelp Botany Building MO90, 6009 Crawley, WA, Australia reaching a maximum length of about 2 m. It has its main E-mail: [email protected] distribution in the southern hemisphere where it is Tel.: +61-8-93802246 Fax: +61-8-93801001 common, often dominant, in the warm-temperate parts of South Africa (Field et al. 1980), Australia (Steinberg M. Coleman and Kendrick 1999) and New Zealand (Choat and Schiel Centre for Research on Ecological Impacts of Coastal Cities, University of Sydney, Marine Ecology Labs (A11), 2006 Sydney, 1982). E. radiata is also found in the northern hemi- NSW, Australia sphere, but so far it has been described only from Oman on the Arabian Peninsula (Bolton and Anderson 1994). A. Fairhead Department of Environmental Biology, University of Adelaide, It is likely, however, that a revision of the genus Ecklonia North Tce, 5005 Adelaide, SA, Australia would include one North African and several Japanese Ecklonia species under E. radiata (Bolton and Anderson S. Miller Department of Botany, University of Otago, PO Box 56, Dunedin, 1994), increasing its northern hemisphere distribution New Zealand considerably. M. Thomsen E. radiata is remarkably successful in terms of its Department of Environmental Science, University of Virginia, ecological dominance and it is probably the most Clark Hall, Charlottesville, VA 22903, USA abundant macroalga in Australia (Steinberg and Kend- 48 rick 1999). It has a profound influence on associated geographic range of Ecklonia radiata from its northern limit in algal assemblages (Kendrick et al. 1999; Melville and Western Australia (Kalbarri: Womersley 1987) and south around Australia to Sydney, and New Zealand. Three random sites were Connell 2001) because canopies of large macroalgae sampled within each location. Large, presumably fully developed, modify the physical environment (Eckman et al. 1989; individuals were targeted during the sampling (Rice et al. 1985; Kennelly 1989; Critchley et al. 1990). The contribution Jackelman and Bolton 1990). Thus, in situ a SCUBA diver hap- of individual kelps to modification of the environment is hazardly collected large stage 3 sporophytes (sensu Kirkman 1981) a function of their morphology (Velimirov and Griffiths by carefully dislodging the holdfast from the reef with a dive knife. Only solitary individuals where the holdfasts did not overlap were 1979; Kennelly 1989). Consequently, if E. radiata ex- collected, and collections were generally made within an area of a hibits systematic spatial variation in morphology, it will few square metres. The collected specimens were kept moist and likewise systematically exert spatially variable effects on refrigerated until processing. algal assemblages. This may be a factor contributing to Within each site 11 morphological characters were sampled from n=5–10 individuals (Table 1). These morphological charac- the large variability commonly encountered in compar- ters were chosen to cover most parts of the thallus (see illustrations isons of E. radiata assemblages across various spatial in Bolton and Anderson 1994). The morphological characters and scales (Kennelly and Underwood 1992; Kendrick et al. procedures for measurement are given in Table 2. 1999; Fowler-Walker and Connell 2002). The objective of the present study was to quantify the morphological variation in E. radiata along its geo- Data analyses graphic range in Australasia. Geographic separation Firstly, we tested whether the morphology of Ecklonia radiata is may cause morphological differences among populations different among locations along its geographic range in Australasia of macroalgae as genetic exchange among populations (H0: there are no differences in E. radiata morphology among lo- becomes restricted (Kusumo and Druehl 2000) and they cations separated by 10s–1,000s of kilometres). Each morphologi- are exposed to grossly different environmental condi- cal character was tested with a one-way ANOVA (Zar 1996) where sites were nested within locations. Before testing, the data set was tions. These differences may be continuously changing balanced by randomly removing individuals from each of the sites following major environmental clines in, for example, to meet the lowest common denominator (n=5; cf. Table 1) and insolation, temperature or salinity (Russell 1986; Kalvas checked for homogeneity of variances (Cochran’s C-test: Winer and Kautsky 1998). Consequently, we tested the hy- et al. 1991). Because of the large number of tests (11 ANOVAs) Bonferroni corrected a-values (a/11) were used to determine sig- potheses that E. radiata has different morphology at nificance levels. Further, the total variation was decomposed and different locations and that the degree of morphological partitioned into the relative contribution from each level of nesting, difference depends on spatial distances among locations. and the relative magnitude of effect (x2) was calculated (Graham and Edwards 2001). Multivariate analyses were performed in PRIMER v5 (Clarke and Gorley 2001). To test the independence of each character, Materials and methods product moment correlation was performed among all pairs of variables using the ‘‘Draftmans plot’’ routine. A normalised eu- Sampling clidean distance dissimilarity matrix was calculated from untrans- formed data using the full unbalanced data set on all Sampling was carried out during summer from November 2000 to morphological characters. Difference in morphology among loca- March 2001. Eleven locations were selected; eight along the south- tions was tested with a two-way nested ANOSIM (analysis of west Australian coastline and three from other Australasian regions similarities) with sites nested in locations, followed by pairwise (Fig. 1; Table 1). Locations within Western Australia were selected comparisons. The data were represented graphically in non-metric on the basis of accessibility and proximity. Locations outside multidimensional scaling (nMDS) plots. SIMPER (similarity of Western Australia were found by requesting collaboration via the percentages) was performed to identify which morphological ALGAL-L email list. These 11 locations span 18° north–south and characters contributed most to the observed patterns. 37° east–west, and they cover >6,000 km of coastline including the Secondly, we tested whether the magnitude of morphological difference was a function of distance between locations (H0: there is no correlation between distance and the degree of morphological similarity among locations). The matrix of pairwise R-values from the ANOSIM was correlated (Spearman rank correlation) to an among-location distance matrix generated from GPS (global po- sitioning system) co-ordinates (cf. Table 1). Results Spatial variability of individual morphological characters Most morphological characters of Ecklonia radiata showed considerable variation among locations (Ta- bles 3, 4). For example, total thallus length differed by Fig. 1 The geographical position of sampling locations. Global >65 cm between the individuals
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