The influence of habitat structure on nearshore fish assemblages in a southern Australian embayment: Comparison of shallow seagrass, reef-algal and unvegetated sand habitats, with emphasis on their importance to recruitment

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Abstract

Assemblage structure and juvenile recruitment of fishes was compared amongst three habitats: seagrass, Heterozostera tasmanica (Martens ex Aschers.) den Hartog; reef-algal; and unvegetated sand. Sampling was conducted monthly from October 1993 to March 1994 at three locations in Port Phillip Bay, southern Australia. A fine-mesh seine net was used to sample subtidally at a depth of approximately 0.5 m. Fish assemblages differed primarily between structured habitats and unvegetated sand, and a number of species previously reported to occur in seagrass habitat were also found to utilise reef-algal habitat. Species richness was highest in seagrass and lowest in unvegetated sand with significant differences amongst all habitats. Total abundance varied between habitats depending on location and month examined, but the most common pattern was highest abundance in seagrass and lowest in unvegetated sand, with a significant difference only between seagrass and unvegetated sand. At the level of individual taxa, pipefishes of the genus Stigmatopora showed a strong preference for seagrass habitat from an early juvenile stage. The King George whiting, Sillaginodes punctata (Cuvier and Valenciennes), showed a complex relationship with habitat, occurring on both seagrass and reef-algae immediately after settlement, but, with growth, showing an increasing preference for reef-algae before finally shifting to unvegetated sand approximately 4 months after settlement. Other species previously found as juveniles on seagrass beds also recruited to reef-algal habitats. Strong locality effects were also found, particularly for King George whiting. This variation was unlikely to be related to habitat structure, because macrophyte biomass showed much greater variation within locations than amongst locations. We conclude that while the presence of structure per se is sufficient for the recruitment of many species, some taxa will discriminate amongst habitats based on structural characteristics.

Introduction

There is wide acceptance that seagrass beds are important `nursery' areas for fishes (Pollard, 1984, Bell and Pollard, 1989), although there are exceptions where seagrasses do not appear to act as nursery areas (Bell and Harmelin-Vivien, 1982, Heck Jr and Thoman, 1984). Reefs and associated algae, and unvegetated sediments, have also been reported as important nursery areas, albeit much less frequently than is the case with seagrass. In the Mediterranean, juvenile fish were mainly associated with shallow, sublittoral reefs, rather than deeper seagrass (Posidonia) beds (Bell and Harmelin-Vivien, 1982). Macrophyte (algae and seagrass) debris in surf zones apparently forms an important nursery habitat for fishes in Western Australia (Robertson and Lenanton, 1984). Sogard and Able (1991)found that seagrass, Zostera, and algae, Ulva, were superior habitats (based on abundances) compared with adjacent unvegetated habitat. Moderately exposed beaches in South Africa may provide a nursery habitat as important as estuaries (Bennett, 1989). Finally, unvegetated sediment adjacent to seagrass, in contrast to the same habitat some distance (greater than 100 m) from seagrass, was an important habitat for juveniles of a number of commercial species in New South Wales (Ferrell and Bell, 1991). Thus, it is possible that the importance of seagrasses may have been over-emphasised relative to other habitats in terms of their nursery function.

The emphasis on the importance of seagrass over other habitats may partly relate to the dominance of this habitat in areas of larval settlement. Bell et al. (1987)conducted experiments on settlement to artificial seagrass that included cage treatments. Results showed high settlement to cage controls with no associated artificial seagrass, leading to the suggestion that shelter per se was critical, rather than the particular type of shelter. In New South Wales, seagrasses usually occur in the middle reaches of estuaries, while rocky reef dominates at the entrance and mangroves dominate in the upper brackish reaches (Bell and Pollard, 1989). Thus, emphasis on the importance of seagrass for certain species of juvenile fish may simply reflect a tendency for larvae of these species to settle in the middle reaches of estuaries (Bell and Pollard, 1989).

The present study compares the diversity and abundance of fishes, including the recruits and juveniles of commercial species, amongst shallow (<1 m) reef-algal, seagrass and unvegetated sand habitats in Port Phillip Bay, Australia. In Port Phillip Bay, reef, seagrass and sand habitats can be found interspersed at the same location, allowing a test of whether the presence of structure per se is more important than the specific structural characteristics. The study was conducted at locations known to span a wide range of fish-recruitment levels, and over the time period of highest fish recruitment (Jenkins et al., 1993, Jenkins et al., 1997b).

Section snippets

Study area

Port Phillip Bay is a large, semi-enclosed, predominantly tidal embayment linked to the ocean of Bass Strait by a narrow entrance (Fig. 1). The hydrodynamics are characterised by: an entrance region, where fast (3 m s−1) ebb and flood jets dominate the circulation; a large flood-tidal delta, known as the Sands region, where strong currents occur in the major channels; and an `inner' zone, where tidal currents are weak (Black et al., 1993). On the western side of Port Phillip Bay, tidal currents

Fish assemblages

Differences were apparent in the assemblages associated with each habitat (Table 1Table 2Table 3). Pipefishes of the genus Stigmatopora were the dominant taxon of fishes collected, occurring primarily in seagrass habitat. A number of groups such as the odacids, clinids, and monacanthids were common on both reef-algae and seagrass but not on unvegetated sand. In contrast, pleuronectids and leptoscopids were restricted to unvegetated sand habitats. Some groups, such as the atherinids, were

Discussion

The hypothesis that the presence of structure per se is more important than the characteristics of the habitat (Bell et al., 1987) is supported to the extent that juveniles of a number of species previously found recruiting to subtidal seagrass, including leatherjackets (Monacanthidae) and King George whiting (Sillaginidae) (Robertson, 1977Jenkins et al., 1997bEdgar and Shaw, 1995), were also common on reef-algae in this study. The greatest difference amongst fish assemblages was between the

Conclusions

In conclusion, seagrass and reef-algal habitats had similar assemblages of fish with the exception of pipefish, Stigmatopora spp., that were dominant in seagrass, and Sillaginodes punctata, that showed a preference for reef-algal habitat over seagrass a month or two after settlement. Unvegetated sand habitats, however, had quite a different assemblage from the structured habitats. Species richness and abundance were generally highest in seagrass, intermediate in reef-algae and lowest on

Acknowledgements

We wish to thank R. Watson, M. Holloway and A. Bellgrove for assistance with field sampling. Comments on drafts of the manuscript from G. Edgar, G. Watson, R. Connolly, P. Hamer and D. Welsford were much appreciated. Funding for this project was provided by the Fisheries Research and Development Corporation and the Australian Research Council.

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