Marine Environmental Research 47 (1999) 373387

Do urban structures inuence local abundance and diversity of subtidal epibiota? A case study from Sydney Harbour, Australia
S.D. Connell*, T.M. Glasby
Centre for Research on Ecological Impacts of Coastal Cities, Marine Ecology Laboratories A11, University of Sydney, Sydney, NSW 2006, Australia Received 6 April 1998; received in revised form 14 September 1998; accepted 12 October 1998

Abstract In an age when human modication of natural substrata is increasingly cited as an agent of population decline and extinction, understanding the role of articial surfaces as surrogate habitats for natural surfaces is critical. It has been predicted that the addition of new habitats to an area can lead to increases in species abundance and diversity. We tested this hypothesis by contrasting assemblages of subtidal epibiota on natural reef and six common urban surfaces in Sydney Harbour, Australia. All surfaces were in shallow water and consisted of rocky reef, sandstone (brick) retaining walls, breglass and concrete pontoons, concrete pilings and wooden pilings with bark and stripped of bark. Assemblages of epibiota on sandstone surfaces (natural rocky reefs and sandstone retaining walls) diered from non-sandstone surfaces. The major distinguishing features of sandstone surfaces were the large cover of coralline algae and small number of taxa. Assemblages on pilings and pontoons were most dierent from those on sandstone surfaces and relatively similar to each other. There were, however, some dierences which seemed to be consistent with features such as type of surface (concrete vs wood) and arrangement of surface (oating pontoons vs xed pilings). We suggest that articial structures may increase the abundance and diversity of subtidal epibiota in the shallow areas of an estuary, but are not surrogate surfaces for epibiotic assemblages that occur on nearby natural rock. It would appear that urbanisation of estuarine habitats has consequences for the identity, diversity and abundance of subtidal epibiota. # 1999 Elsevier Science Ltd. All rights reserved.
Keywords: Fouling organisms; Sessile; Articial habitats; Biodiversity; Disturbance

* Corresponding author. Tel.: +61-2-9351-4933; fax: +61-2-9351-6713; e-mail: sconnell@zoology. adelaide.edu.au 0141-1136/99/$see front matter # 1999 Elsevier Science Ltd. All rights reserved. PII: S014 1-1136(98)0012 6-3

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1. Introduction Habitat structure has long been considered an important determinant of the number, identity and abundances of species present in biological communities (Abele, 1974; Heck, 1979; Jones, 1991; MacArthur, MacArthur, & Preer, 1962; Menge & Sutherland, 1976; Williams, 1943). One particular feature of habitat structure is the material from which the habitat is made, i.e. the type of substratum. In an age when human modication of natural substrata is increasingly cited as an agent of population decline and extinction, understanding the role of articial surfaces as surrogate habitats for natural surfaces is critical. In estuaries around Sydney, rocky reefs provide the predominant natural habitat for marine plants and animals that attach to subtidal hard substrata. These natural habitats, however, continue to be destroyed and replaced with man-made structures, particularly vertical surfaces (e.g. sandstone walls and pilings), which are made from dierent substrata (e.g. concrete, breglass, wood). Notwithstanding the wide use of articial surfaces to investigate fouling assemblages (see review in Cairns, 1982), few studies have specically compared articial and natural surfaces for the purposes of understanding their relative eects on species diversity and abundance (but see McGuinness, 1989). Many studies have investigated the population dynamics of epibiota in the marine environment. Traditionally, a variety of types of articial surfaces have been used as surfaces for settlement because they are manipulable and easy to use. The results of these studies suggest that dierent substrata can be expected to aect settlement (Crisp & Ryland, 1960; Cuomo, 1985; Russ, 1977; Walters & Wethey, 1996) and subsequent development of intertidal and subtidal assemblages (Anderson & Underwood, 1997; Butler, 1991; Keough, 1984; McGuinness, 1989; Walters & Wethey, 1996). Despite evidence of the potential importance of substratum type to sessile assemblages, few studies have specically addressed the eect of urban structures on subtidal epibiota (Glasby, in press). In this study, we tested the prediction that assemblages of epibiota associated with urban structures of particular material dier and that these are distinct from assemblages on the natural reef. A noteworthy feature that characterises an assemblage is the number of constituent organisms, i.e. species diversity. Species diversity has become a major issue in conservation biology (Chapin et al., 1998; Tilman et al., 1997), particularly in conjunction with the modication of habitat (Dunning, Danielson, & Pulliam, 1992). It has been suggested that the creation of new habitats through urbanisation not only increases habitat diversity, but this in turn causes an increase in species diversity (Rebele, 1994). There has, however, been no direct test of this in the marine environment. The idea that habitat diversity promotes species diversity was recognised as early as the 1940s (Williams, 1943). Subsequently, it has been demonstrated that increases in the diversity of habitats can increase the number of species in an assemblage (Bohninggaese, 1997; Douglas & Lake, 1994). Yet in some instances, an increase in habitat diversity has actually decreased the number of species in an assemblage and/or the abundance of individuals (Heck, 1979; McGuinness & Underwood, 1986). Hence, we also tested whether species diversity (number of taxa)

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of subtidal epibiota growing on vertical surfaces was enhanced by the presence of articial surfaces. This study was designed to compare the number of taxa, types of organisms and abundance of individuals of subtidal epibiota among rocky reefs and common articial surfaces in Sydney Harbour, Australia. At least six types of articial surfaces are common in shallow subtidal areas of Sydney harbour: breglass pontoons, concrete pontoons, wooden pilings with bark, wooden pilings stripped of bark, concrete pilings and articial sandstone retaining walls. The major surfaces of most of these structures are orientated vertically, hence we compared the vertical surfaces of these six structures to the vertical surfaces of the predominant natural hard surface, sandstone rocky reef. 2. Methods 2.1. Sites and sampling methods Assemblages of subtidal epibiota were studied in December 1997 (summer) at two sites in Middle Harbour, the northern part of Sydney Harbour, Australia (34 01H S, 151 11H E, Fig. 1). Epibiota were dened as assemblages of marine algae and sessile invertebrates growing on hard substrata. The vertical surfaces of seven types of substrata were sampled: natural rocky reef (sandstone), articial sandstone retaining walls (bricks of sandstone), wooden pilings with bark surfaces, wooden pilings stripped of bark, concrete pilings, concrete pontoons and breglass pontoons. Pontoons ($2.53.50.4 m deep) were moored to concrete pilings which together act as platforms and anchorage for boating activities. Sampling involved photographing surfaces (1523 cm) at a depth of 0.2 m below Mean Low Water Springs. Five replicate photographs were taken of each type of surface; these represented dierent and haphazardly chosen structures. This was repeated at each of two sites (Fig. 1). Percentage covers of taxa were estimated from projected photographs in which the identities of taxa were recorded under 40 evenly spaced points across a 520 cm transect placed haphazardly on the image. Any taxa in quadrats not sampled under the points were noted as occurring in the sample. A pilot study indicated that the size and replication of transects, and number of points used to estimate percentage covers of taxa generally gave relatively precise estimates of abundance (unpublished data). 2.2. Analytical methods A visual assessment of the dissimilarities in the structure of assemblages among surfaces was provided by a dendrogram; dissimilarities among surfaces and between sites were provided by a non-metric multi-dimensional scaling ordination (nMDS). The centroids used in the nMDS plot used the average of all replicates per surface for each site (n=5). The Bray-Curtis similarity measure was used to calculate similarities among replicate observations (Bray & Curtis, 1957) which were fourth-root

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Fig. 1. Map showing the two study sites within Middle Harbour, Sydney Harbour.

transformed. Two one-way analyses of similarities (ANOSIM) were done (Clarke, 1993) on all replicates to test for dierences among surfaces and between sites. The signicance level was adjusted for multiple comparisons associated with pairwise tests (Bonferroni procedure; Rice, 1989); a=0.002 for comparisons of surfaces and 0.007 for `Site'. Similarity percentages (SIMPER) were used to determine which taxa primarily (i.e. accounted for 80% of the similarity) contributed to average similarity within a surface and to provide measures of the relative dissimilarity among surfaces (Clarke, 1993). Multivariate analyses were done using a suite of multivariate techniques included in the PRIMER program (Plymouth Routines In Multivariate Ecological Research; Clarke, 1993). Taxa that were present in more than 15% of the 70 photographs were analysed with analysis of variance (ANOVA) according to Winer, Brown, and Michels (1991) and Underwood (1981). Two-factor ANOVA treated `Surface' as xed and `Site' as random and orthogonal to `Surface'. Post-hoc pooling was used to provide a more powerful test of `Surface' when the interaction `SurfaceSite' was nonsignicant (p>0.25; Winer, Brown, & Michels, 1991). Prior to analysis, data were tested for homogeneity of variances using Cochran's C-Test (Underwood, 1981).

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Heterogeneous data were Arc-sine transformed and if this transformation did not remove heteroscedasticity, the raw data were used for the analysis with the more conservative probability of 0.01. 3. Results 3.1. Comparisons of assemblage structure among surfaces Natural rocky reefs and in particular sandstone retaining walls (both sandstone) supported the most distinct assemblages (Figs. 2 and 3). Correspondingly, measures of dissimilarity indicated that assemblages on sandstones surfaces were most dierent from the other surfaces (Table 1), and pairwise tests indicated that the two sandstone structures (rocky reefs and sandstone walls) did not dier from each other (ANOSIM: p>0.002), but diered from the other surfaces (ANOSIM: p<0.002). Of the remaining surfaces, assemblages on breglass pontoons were the most distinctive (Figs. 2 and 3, Table 1). Although assemblages on concrete pontoons were most similar to breglass pontoons (Table 1), pairwise tests revealed that

Fig. 2. Dendrogram showing the relative dissimilarities of assemblage structure among the seven surfaces.

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Fig. 3. Multi-dimensional scaling ordination showing the relationship between surfaces and sites. ~=sandstone retaining walls, &=rocky reef, !=wooden striped-pilings, +=concrete pilings, =concrete pontoons, ^=wooden barked-pilings, *=breglass pontoons. The symbols in lighter ink refer to Site 1, those in heavier ink to Site 2. Table 1 Bray-Curtis measures of dissimilarities between surfaces (the larger the value the more dissimilar the assemblage) Surface Fibreglass pontoon Concrete pontoon Concrete piling Bark piling Stripped piling Natural reef Sandstone wall Surface Measure of dissimilarity 46.7 58.7 55.9 56.7 69.7 79.9 F. pontoon

42.3NS 45.3NS 43.5NS 65.5 81.4 C. pontoon

43.5NS 46.7NS 65.3 80.4 C. piling

50.8 70.6 84.3 B. piling

66.1 82.1 S. piling

50.9NS Natural reef

All pairwise comparisons are signicantly dierent (p<0.002) unless indicated NS.

assemblages diered signicantly between these surfaces (ANOSIM: p<0.002). Other than sandstone substrata, surfaces with the most similar assemblage structure were concrete pilings and concrete pontoons (Table 1). Assemblages on these concrete substrata did not dier from those on wooden pilings (barked and unbarked),

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but assemblages on the two types of wooden pilings diered from each other (ANOSIM: p<0.002). In general, spatial variability (between sites) did not appear to have a greater eect than surface-type on assemblage structure (Fig. 3). No signicant dierences were detected between sites for any surface, except possibly sandstone retaining walls. There was, however, evidence to suggest that the composition of assemblages on each surface was somewhat dependent on site; points representing Site 1 (Fig. 3) are further up the plot than points from the equivalent surfaces at Site 2. For six of the seven surfaces there were no signicant dierences in the composition of assemblages between the two sites (p>0.02). The Bonferroni correction for multiple comparisons reduced a from 0.05 to 0.007, but the smallest possible level of probability for pairwise comparisons between sites (n=5) was p=0.008. This signicance level occurred for the comparison between sites for assemblages on sandstone retaining walls. Thus, it is possible that the sandstone wall sites diered and this was supported by the large R value (0.656) for that comparison. 3.2. Comparisons of individual taxa among surfaces Of 31 taxa sampled across all surfaces and sites (Appendix A), 21 are known to occur on natural reef in Middle Harbour; 14 were detected in this study and 7 others in Glasby (1997). Hence, articial surfaces supported 10 species that have not been sampled on rocky reefs. Importantly, no one articial surface supported the same suite of species that was typical of rocky reefs. Univariate analyses indicated that there were signicantly more taxa on breglass pontoons than sandstone retaining walls and that there were no dierences in the number of taxa among other surfaces (Fig. 4a, Table 3a, SNK test). In general, the cover of no taxon varied greatly among the articial surfaces (breglass, concrete, wood), but some taxa varied greatly between these and sandstone surfaces (natural reef and sandstone walls). A major distinguishing feature of sandstone surfaces was the domination of space by coralline algae, particularly on sandstone retaining walls (Fig. 4b, Tables 2 and 3b). These algae almost completely covered sandstone retaining walls and were very abundant on natural rock, but were absent or uncommon on the other ve surfaces. This extensive cover of coralline algae was associated with reduced abundance of other taxa, particularly lamentous algae and serpulid polychaetes (Hydroides spp.; Fig. 4c,d, Table 3b,d). Encrusting bryozoans (primarily Watersipora subtorquata) were abundant on breglass and concrete pontoons. Fibreglass pontoons supported the greatest abundance of bryozoans at one site (Site 2, Table 3e). The SNK test was unable to identify dierences among surfaces for Site 1, but there was a trend for a greater cover of bryozoans on the two types of pontoons and natural reef than any other surface (Fig. 4e). Most notably, no bryozoans occurred on sandstone retaining walls at either site (Fig. 4e). Despite the similarities in the coverage of taxa among the three types of piling, pilings without bark where notable for the percentage cover of unoccupied (bare) space (Fig. 4f, Table 3f). Spirorbid polychaetes, which are comparatively small

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Fig. 4. Graphs of taxa showing the mean percentage cover among surfaces and between sites. Shaded bars represent Site 1 and unshaded Site 2. FPo, breglass pontoons; CPo, concrete pontoons; CPi, concrete pilings; BPi, bark pilings; SPi, stripped pilings; NR, natural rock; SW, sandstone retaining walls.

Table 2 Results of SIMPER analysis showing the taxa that primarily (80%) contributed to average similarity within a surface, i.e. `Characteristic taxa' Surface Natural rock Sandstone wall Fibreglass pontoon Concrete pontoon Concrete piling Bark piling Stripped piling Characteristic taxa Coralline algae Coralline algae Green algae, Bryozoan (Waterspoira subtorquata) Hydroides, spirorbids, barnacles Hydroides, spirorbids Hydroides, bare spacea Hydroides, red lamentous, barnacles (Site 1), sponge sp. (Site 2)

Green/brown lamentous algae was a taxon characteristic of all surfaces, except sandstone retaining walls (not listed). a Bare space was included in SIMPER analyses, but was not counted as total or novel taxa.

organisms, contributed to a great deal of the total cover on pilings and natural rock, but very little to the cover on sandstone walls and breglass pontoons (Fig. 4g, Table 3g). Although signicant dierences among surfaces were detected

S.D. Connell, T.M. Glasby/Marine Environmental Research 47 (1999) 373387 Table 3 ANOVA comparing the percentage cover of selected taxa among seven surfaces at two sites Source Surface Site SuSi Residual Surface Site SuSi Residual Surface Site SuSi Residual df 6 1 6 56 6 1 6 56 6 1 6 56 MS F p MS F p MS F p

381

(a) Number of taxa * 25.88 7.42a 0.23 0.07 4.33 1.25 >0.25 3.56 (d) Hydroides 741.42 6.62 5.04 0.08 111.91 1.84 61.00 * NS

(b) Coralline algae 11895.42 43.41 *** 165.09 1.20 274.05 1.99 NS 137.63 (e) Bryozoans 742.49 4.00 0.75 0.01 185.47 3.22 57.67 NS **

(c) Filamentous algae 4793.63 7.92 * 2035.80 6.65 605.60 1.98 NS 306.21 (f) Bare 157.44 2.23 19.11 36.38 space 8.24 0.06 0.53 * >0.25

(g) Spirorbids 158.67 3.90a * 78.80 1.98 48.84 1.23 >0.25 39.84

(h) Porifera 39.57 2.38 * 0.36 0.02 12.44 0.73 >0.25 17.14

(i) Ceramiales 292.09 1.09 272.10 2.44 266.77 2.39 111.53

NS *

Post-hoc pooling of the interaction term with the residual enabled a more powerful test of the main factor `Surface' (p>0.25; Winer, Brown, & Michels, 1991). NS, p>0.05; *p<0.05; **p<0.01; ***p<0.001. The critical value of `a' was adjusted to allow for signicant heterogeneity of variances (Cochran's C-test p<0.05). The comparison of sites was not relevant for testing the hypothesis. Transformations: untransformed=a, b, c, f, h, i; Arc-sine=d, e, g. a F-ratios aected by pooling and resultant values are given.

for porifera and red lamentous ceramalean algae (Fig. 4h,i, Table 3h,i), SNK tests could not detect which means diered. 4. Discussion The results of this work lead to two main conclusions about assemblages of epibiota in shallow depths of estuaries. First, the structure of assemblages on natural surfaces may dier greatly from articial surfaces at similar depths, and any dierences among articial surfaces are likely to be smaller. Second, we suggest that articial structures may increase the abundance and diversity of subtidal epibiota in the shallow areas of an estuary, but are not surrogate surfaces for epibiotic assemblages that occur on nearby natural rock. 4.1. Eects of substratum on assemblage structure The assemblages of epibiota on natural rock (sandstone) and sandstone retaining walls diered from those on the wooden, concrete and breglass surfaces of pilings and pontoons. The major distinguishing features of sandstone surfaces were the dierence in identity of species in the assemblage and the abundance of individual

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taxa. This appeared to be driven primarily by the domination of space by coralline algae which correspondingly reduced the abundance of other taxa. Note that it is possible that other taxa occurred under the thick mat of coralline algae. The assemblages among pilings and pontoons (not sandstone) showed some striking similarities. The two structures with the most similar assemblages were concrete pilings and concrete pontoons. A possible explanation for this is that concrete has certain physical or chemical properties which aect the settlement, growth or survival of organisms. For example, the high alkalinity at the surface of concrete, due to the leaching of calcium hydroxide from cement, enhances the settlement of oysters under laboratory and eld conditions (Anderson, 1996). While strippedpilings and barked pilings were both made of wood, their physical and chemical properties are also likely to vary, and this may explain why the assemblages on these two surfaces diered. Pilings with bark are notable by the large amount of bare space (up to 35%) and this may be due to features of the bark that reduce settlement and subsequent recruitment of some taxa. This seems likely given that the bark is left on pilings to help prevent deterioration of the wood by fouling organisms and boring molluscs. Dierent types of structures are also likely to be correlated with topographic complexity and area of surface. Previous work has shown that complex surfaces can provide sites for settlement and improved opportunities for attachment, growth and survivorship of organisms (Walters & Wethey, 1996). Moreover, it is well known that the number of species within a habitat often increases with an increase in the size of habitat (Gleason, 1922; McGuinness, 1984; Simberlo & Abele, 1982) and this can happen at relatively small scales for subtidal epibiota (Butler, 1991; Keough, 1984). Thus, dierences in size (and shape) of a structure could be partially responsible for the observed dierences in assemblages on these surfaces. Although the characteristics of the surface appear to be an important determinant in structure of assemblages, the arrangement of the surface may also be inuential. While breglass pontoons supported the most distinct assemblages of non-sandstone surfaces, concrete pontoons supported the least dissimilar assemblage to breglass pontoons. This indicates that although breglass and concrete surfaces have very dierent eects on assemblage structure, pontoons themselves have very distinctive eects on assemblage structure. A unique aspect of pontoons is that they oat and attached organisms are not subjected to changes in depth associated with the rise and fall of the tide. No studies have compared the development of assemblages with and without the inuence of tides, but it is likely that constant depth and characteristics associated with depth (e.g. light) have considerable eects on the development of assemblages. The feeding activities of sh and invertebrates have been shown to have profound eects on assemblages of subtidal epibiota (Breitburg, 1985; Russ, 1980). Moreover, it is widely speculated that predatory sh are a major structuring force on patterns of abundance of subtidal epibiota (Harris & Irons, 1982; Sebens, 1986; Walters & Wethey, 1996; Young & Chia, 1984) and also contribute to dierences in assemblages of intertidal epibiota growing on plates constructed of various substrata

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(McGuinness, 1989). In Sydney Harbour, there is substantial variability in the abundance and composition of sh around pontoons, pilings and natural reef separated by only tens of metres (Connell, personal observation) and sh are major predators of oysters on pilings (Connell, unpublished data). Hence, grazing and predation particularly by sh cannot be discounted as a factor contributing to differences in assemblages of epibiota among such structures. Finally, variation in the age of the various surfaces may account for a large proportion of the dierences observed between natural reef and articial surfaces. Urban structures are considerably younger than rocky reef and are likely to have been introduced at dierent times. Variation in age is an intrinsic property of urban structures. Consequently, many structures would have a unique history of various events of recruitment, mortality and succession. It is well known that assemblages can be aected by temporal heterogeneity (Menge & Sutherland, 1976) associated with the timing of biological or physical disturbance (Breitburg, 1985; Dean & Hurd, 1980), and stage of succession (Connell & Slatyer, 1977) or time since disturbance (Anderson & Underwood, 1994). Hence, temporal heterogeneity will be an important source of variation in the structure of assemblages between natural reef and articial surfaces, especially since the addition of urban structures to marine habitats has and will continue to occur haphazardly through time. 4.2. Habitat diversity and species diversity The idea that the addition of new habitats can increase species diversity is not new to ecology (Connor & McCoy, 1979; Williams, 1943), and has been adapted as an important theme in the subdisciplines of conservation biology (Barbault, 1995; Kaplan, 1993; Walting, 1997) and urban ecology (Rebele, 1994). It has been recognised that urban structures are habitats for a variety of plants and animals in terrestrial systems (Rebele, 1994), but until now this has not been studied in marine systems. We were unable to refute the model that the addition of urban structures has resulted in a greater number of species of attached plants and animals at shallow depths. Although it is clear that urban structures modify the identities and numbers of species within an area, the idea that this has a positive eect on species diversity needs to be treated with extreme caution (Glasby & Connell, in press). The addition of structures to the natural habitats often results in the fragmentation of natural habitats and many studies have demonstrated a decline of diversity in habitats after fragmentation (Andren, 1994; Devries, Denboer, & Vandijk, 1996; Spellerberg, 1991). Furthermore, the addition of exotic species, which enhances species diversity, may have catastrophic eects on the abundance of plants and animals and on the preservation of endangered species (Lodge, 1993). Articial surfaces may be surrogate habitats for some species, but our results suggest that as habitat for assemblages of epibiota, natural reef may not be adequately replaced by urban structures. Importantly, articial surfaces create new habitat. Assemblages of epibiota on them are dierent from those on natural reef at the same depth, thereby altering the distribution and abundances of epibiotic

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organisms. The current xation with species diversity, therefore, may be inappropriate if it diverts attention from understanding the eects on the abundance of the component species. Alteration of habitat may have profound eects on the abundance of particular species and in turn these changes may cause negative or positive impacts on the broader environment. The notion that urban structures may have positive and negative impacts on the environment parallels developments in sheries science. Although articial structures are seen as eective tools to enhance the diversity and productivity of commercially important species (Pickering & Whitmarsh, 1997), it is recognised that they can degrade the environment (Chou, 1997). Consequently, sheries science has identied the need for and begun research on how dierent types of structures aect a shery (Bassett, 1994; Collins, Jensen, Lockwood, & Lockwood, 1994). In a similar context, assessments of how alternative urban structures aect the broader marine environment are required. In conclusion, our results clearly indicate the potential for both the type of surface (e.g. wooden pilings with vs without bark) and the size/shape of the same type of surface (e.g. concrete pontoons and concrete pilings) to aect the composition and abundance of the assemblage. These results taken with others (Anderson & Underwood, 1994; McGuinness, 1989; Michener & Kenny, 1991) highlight that it is not only the identity of species that vary among surfaces, but also the abundance of the component species. Future research may be well directed in establishing the types of species and how their abundances are aected by the addition of urban structures. An increase in the number of urban habitats per se is not necessarily benecial to the environment if the associated changes in species composition and abundance lead to the degradation of other components of estuarine areas. Acknowledgements The Centre for Research on Ecological Impacts of Coastal Cities and this work was supported by an ARC Special Research Centre Grant. We thank G. Chapman, M. Lindegarth and A. Underwood for their advice and J. Cunningham and G. Houseeld for assistance with eldwork and persisting with remarkably average diving.

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Appendix A
Taxa identied on rocky reef and urban structures. `Surface' indicates the surface from which the taxon was sampled: A (articial surfaces), N (natural reef) and B (both articial surfaces and natural reef) Phylum/Class Chlorophyta Taxa Enteromorpha spp. Ulva lactuca Cladophorales Brown/green lamentous complex Ceramiales complex Encrusting unid. Rhodymenia australis Corallina ocinals Peyssonnelia spp. Colpomenia sinuosa Sargassum spp. Zonaria sp. Dictyota dichotoma sp. 1 sp. 1 sp. 2 sp. 3 sp. 4 sp. 5 Spirorbidae Hydroides spp. Balanus trigonus Saccostrea commercialis Mytilus edulis Anomia sp. Watersipora subtorquata Schizoporella errata Fenestrulina mutabilis Bugula neritina Styela plicata Pyura sp. Surface Aa Aa A B B N Aa B N A B A A Aa B A Aa Aa A B B B N Aa Aa B A N Aa B Aa

Rhodophyta

Phaeophyta

Anthozoa Porifera

Polychaeta Cirripedia Bivalvia Bryozoa

Ascidiacea

a Taxa also not detected in a previous study of vertical rocky-reefs in Middle Harbour at 0.5 m below Mean Low Water Springs (Glasby, 1997). Four 1515 cm photographic quadrats were sampled in each of six sites during January 1996.

References
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