Fisheries Management and Ecology, 1999, 6, 401±420 Fishing impacts and the degradation or loss of habitat structure S. J. TURNER,* S. F. THRUSH, J. E. HEWITT, V. J. CUMMINGS & G. FUNNELL National Institute of Water and Atmospheric Research, Hamilton, New Zealand Abstract The wider effects of fishing on marine ecosystems have become the focus of growing concern among scientists, fisheries managers and the fishing industry. The present review examines the role of habitat structure and habitat heterogeneity in marine ecosystems, and the effects of fishing (i.e. trawling and dredging) on these two components of habitat complexity. Three examples from New Zealand and Australia are considered, where available evidence suggests that fishing has been associated with the degradation or loss of habitat structure through the removal of large epibenthic organisms, with concomitant effects on fish species which occupy these habitats. With ever-increasing demands on fish-stocks and the need for sustainable use of fisheries resources, new approaches to fisheries management are needed. Fisheries management needs to address the sustainability of fish-stocks while minimizing the direct and indirect impacts of fishing on other components of the ecosystem. Two long-term management tools for mitigating degradation or loss of habitat structure while maintaining healthy sustainable fisheries which are increasingly considered by fisheries scientists and managers are: (1) protective habitat management, which involves the designation of protected marine and coastal areas which are afforded some level of protection from fishing; and (2) habitat restoration, whereby important habitat and ecological functions are restored following the loss of habitat and/or resources. Nevertheless, the protection of marine and coastal areas, and habitat restoration should not be seen as solutions replacing conventional management approaches, but need to be components of an integrated programme of coastal zone and fisheries management. A number of recent international fisheries agreements have specifically identified the need to provide for habitat protection and restoration to ensure long-term sustainability of fisheries. The protection and restoration of habitat are also common components of fisheries management programs under national fisheries law and policy. k e y w o r d s : fisheries management, fishing, habitat degradation, habitat loss, habitat restoration, habitat structure, protected marine and coastal areas. Introduction The wider effects of fishing on marine ecosystems have become the focus of growing concern *Correspondence and present address: S. J. Turner, Department of Conservation, Northern Regional Office, PO Box 112, Hamilton, New Zealand (e-mail: sturner@doc.govt.nz). ã 1999 Blackwell Science Ltd 401 402 S. J. TURNER ET AL. among scientists, fisheries managers and the fishing industry over the past decade (e.g. Fowler 1989; Hutchings 1990; Northridge 1991; ICES 1992; Jones 1992; Dayton, Thrush, Agardy & Hofman 1995). Fishing activities, such as trawling and dredging for fish and shellfish, where mobile fishing gear is towed across the sea bed, have the capability of altering, removing or destroying the complex, three-dimensional physical structure of benthic habitats by the direct removal of biological (e.g. sponges, hydroids, bryozoans, amphipod tubes, shell aggregates and seagrass) and topographic (e.g. sand depressions and boulders) features. These impacts are in addition to documented effects of fishing on sediment dynamics (e.g. sediment suspension and deposition), sediment chemistry (e.g. alteration of the sediment chemistry and changes in the availability of toxic contaminants) and benthic/pelagic nutrient fluxes (e.g. de Groot 1984; Messieh, Rowell, Peer & Cranford 1991; Black & Parry 1994). The present review examines the role of habitat structure and habitat heterogeneity (see Sebens 1991) in marine ecosystems, as well as the effects of fishing on these two components of habitat complexity. It considers three examples from New Zealand and Australia, where available evidence suggests that fishing has been associated with the degradation or loss of habitat structure through the removal of large epibenthic organisms, with concomitant effects on fish species which occupy these habitats. The different management actions, if any, which have been adopted in each case are reviewed, and where documented, the outcomes in terms of the recovery of habitat structure and fisheries resources are discussed. With ever-increasing demands on fish-stocks and the need for sustainable use of fisheries resources, new approaches to fisheries management are needed. Fisheries management needs to address the sustainability of fish-stocks while minimizing the direct and indirect impacts of fishing on other components of the ecosystem. To date, the extent and effects of fishing on habitat structure have been largely neglected in fisheries management. The development of protective habitat management and habitat restoration are reviewed as means of mitigating for degradation or loss of habitat structure while maintaining healthy sustainable fisheries, and the provisions for these management tools in recent international fisheries agreements and fisheries law and policy at a national level are discussed. Role of habitat structure in marine ecosystems Emergent structures, such as rocky outcrops, boulder shoals, epibenthic reef formations (e.g. coral, sabellid, vermetid and oyster reefs), vegetation (e.g. seagrasses, salt marsh, mangroves, kelp and other macroalgae) as well as other topographic features (e.g. shell, burrows, biogenic structures and depressions), provide heterogeneity and structural complexity in marine benthic environments. The structural framework provided by emergent features constitutes an important organizing feature of many ecological systems and is critical to the functioning of the ecosystem as a whole (Ryder & Kerr 1989; Peters & Cross 1992). Emergent structures represent important habitat for a variety of marine organisms, including a number of commercially and recreationally valuable fisheries species. These structures may provide refuge from predation and competition, as well as physical and chemical stresses, or may represent important food resources and critical nursery or spawning habitat (e.g. Thayer, Stuart, Kenworthy, Ustach & Hall 1978; Robertson & Duke 1987; Rozas & Odum 1988; ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 403 Thayer, Colby & Hettler 1988; Heck & Crowder 1991; Safriel & Ben-Eliahu 1991; Sebens 1991; Fonseca, Kenworthy & Thayer 1992; Gotceitas & Brown 1993; Posey & Ambrose 1994; Auster, Malatesta & LaRosa 1995; Langton, Auster & Schneider 1995; Tupper & Boutilier 1995; Perkins-Visser, Wolcott & Wolcott 1996). In addition, these structures modify the hydrodynamic flow regime near the sea floor, with potentially significant ecological effects on food availability, growth, larval and/or juvenile recruitment and sedimentation (e.g. Fonseca, Fisher, Zieman & Thayer 1982; Fonseca, Zieman, Thayer & Fisher 1983; Eckman 1987; Irlandi & Peterson 1991). Although not always essential for the completion of specific lifehistory stages in terms of representing a `demographic bottle-neck', these structures may have a significant positive influence on survival. Habitats do not exist independently of each other, and a marine benthic system may consist of several habitat types which are all integrally linked through biological, chemical and physical processes to form a complex landscape. Species may utilize this entire landscape during their life-cycle (e.g. Kenworthy, Thayer & Fonseca 1988; Parrish 1989; Baelde 1990; Hoss & Thayer 1993; Szedlmayer & Able 1996). Thus, the integrity of the habitat landscape as a whole may be just as important for fisheries resources as the integrity of a particular habitat structure (Sebens 1991). There is abundant evidence in the ecological literature which demonstrates the importance of habitat structure and heterogeneity in influencing faunal abundance, species richness and species composition of invertebrate and fish communities (e.g. Risk 1972; Heck & Wetstone 1977; Luckhurst & Luckhurst 1978; Carpenter, Miclat, Albaladejo & Corpuz 1981; Lewis 1984; Pollard 1984; Pihl 1986; Choat & Ayling 1987; Roberts & Ormond 1987; Rozas & Odum 1988; Bell & Pollard 1989; Bohnsack 1989, 1991; Sale 1991; Sogard & Able 1991; Walters & Juanes 1993; Levin, Talley & Hewitt 1998; Turner & Kendall in press). It is also widely acknowledged that the size, vitality and spatial distribution of populations of many fisheries species are dependent on the quantity and quality of the habitat, although the ecological relationships which constitute this dependence have not been quantified in most cases (e.g. Nixon 1980; Lindall & Thayer 1982; Boesch & Turner 1984; Fonseca, Kenworthy & Thayer 1988; Karr 1991; Peters & Cross 1992; Hoss & Thayer 1993; Langton, Steneck, Gotceitas, Juanes & Lawton 1996; Auster, Watling & Rieser 1997). If fisheries are to be wisely managed, it is essential that fisheries managers understand the extent to which the harvested species are dependent on habitat, together with the long-term implications of degradation or loss of habitat structure on these species, non-target species and the ecosystem as a whole. Degradation or loss of habitat structure Extensive areas of benthic habitat have been lost or their physical integrity compromised as a result of fishing (e.g. sabellid reefs in the Wadden Sea: Reise 1982; Riesen & Reise 1982; seagrass beds in North Carolina: Fonseca, Thayer, Chester & Foltz 1984; Hsiao, Easley & Johnson 1987; Peterson, Summerson & Fegley 1987; oyster bars in Chesapeake Bay: Rothschild, Ault, Goulletquer & HeÂral 1994; sponge communities in the Gulf of Maine: Auster, Malatesta, Langton, Watling, Valentine, Donaldson, Langton, Shepard & Babb 1996). In many areas, the spatial extent (both individually and in aggregate) and severity of physical ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 404 S. J. TURNER ET AL. disturbance, the potentially slow recovery of impacted ecosystems, as well as the frequency of occurrence over time (heavily fished areas may be impacted many times in a year), directly attributable to fishing far exceed the effects of other disturbance agents (e.g. waves, tidal currents, bioturbation processes, and anthropogenic processes such as dredging and extractive activities). Nevertheless, few studies have considered the effects of fishing on the physical modification of habitat structure and the potential implications for associated faunal communities. Likewise, there is little information concerning how, and to what extent, changes in habitat structure affect fisheries resources and contribute to fisheries declines. Habitat degradation or loss as a result of fishing remains the least understood of the environmental impacts of fishing (Committee on Fisheries, Ocean Studies Board, National Research Council 1994, cited in Auster et al. 1996). Because of the inadequate data which are available to address the specifics of the complex ecological interactions involved, it is rarely possible to predict or quantify the potential loss to fisheries production caused by the degradation or loss of habitat structure. An understanding of the extent of this impact, and its effect on populations of marine organisms (both target and non-target species), is essential if fisheries are to be strategically managed by setting appropriate levels of fishing effort which will maximize fisheries production, and therefore, yield. The extent of fishing activities is potentially extremely large, often leaving few, if any, undisturbed habitats within impacted areas. As well as effects on habitat structure per se, there are important implications on a larger scale with regard to effects on habitat heterogeneity, or the patchiness of habitats, across a landscape. Reductions in heterogeneity over large spatial and temporal scales have implications for the maintenance of diversity and stability at the population, community and ecosystem level (Thrush, Hewitt, Cummings & Dayton 1995). As a landscape becomes fragmented, with reduced physical structure occurring between habitat units, movement of adults, recruits and larvae between habitats becomes impeded. Furthermore, as more and more of the habitat structure and the associated communities are destroyed, sources of larvae recruiting into disturbed areas from adjacent habitats will decline, and the recovery-time for biological communities (both habitat-forming species as well as associated species) in these impacted areas will progressively increase. Fishing impacts and the degradation or loss of habitat: examples Three examples from New Zealand and Australia were considered, where available evidence suggested that fishing has been associated with the degradation or loss of habitat structure through the removal of large epibenthic organisms, with concomitant effects on fish species which occupy these habitats. The management strategies (if any) which were adopted in each example are also reviewed. 1. Benthic epifauna and finfish stocks: Tasman Bay, New Zealand Around the northern coast of New Zealand's South Island (Tasman Bay), fishermen have long reported an association between juveniles of a number of commercially important species of fish [including snapper, Chrysophrys auratus (Bloch & Schneider), tarakihi, Cheilodactylus ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 405 macropterus (Bloch & Schneider), and John Dory, Zeus faber (L.)] with clumps and mounds of `coral-like' bryozoans. Bryozoans [predominantly the endemic species Celleporaria agglutinans (Hutton) and Hippomenella vellicata (Hutton)], along with other calcareous frame-building species (e.g. encrusting bryozoans, serpulids and foraminiferans) and associated epifauna (e.g. hydroids, sponges, solitary and colonial ascidians, bivalves, polychaetes, crustaceans, and ophuiroids), form abundant and extensive `coral-like' growths (attaining spans of 0.3 m and heights of 0.15±0.5 m) which provide an important source of shelter and food for juveniles of several fish species (Bradstock & Gordon 1983). With the advent of commercial trawling in Tasman Bay in the 1940s, there was extensive destruction of the bryozoan growths. The size and extent of these important nursery grounds have been progressively reduced, especially since the introduction of synthetic trawl nets (which are less prone to damage by the bryozoan growths) in the 1960s and 1970s (Saxton 1980; Bradstock & Gordon 1983). In some areas, these grounds were virtually destroyed by the late 1970s. Observations in 1980 revealed that, where the growths persisted, these were markedly reduced in size (not exceeding 0.15 m in height above the sea floor) and density (clumps were often several metres apart) (Bradstock & Gordon 1983). There was a marked reduction in the numbers of juvenile fish associated with this decline in habitat structure. Management action In 1980, a region delimiting an area of bryozoan growths was closed to power fishing methods (i.e. trawling, Danish seining and dredging) (Mace 1981). The expectation was that the closure of the fishing grounds would allow the bryozoan growths to regenerate, thereby restoring the fish habitat to its original state and allowing recovery of the fish-stocks. To date, the area remains closed, and according to the Ministry of Agriculture and Fisheries, the closure is permanent. Extraordinarily, habitat recovery has not been monitored (Gordon 1994) and the closure of the fishing grounds has not been accompanied by any scientific research. There is no information with which to assess whether closure of the fishing grounds has led to recovery of the bryozoan growths, and whether fish-stocks are further declining, being maintained or are increasing. 2. Benthic epifauna and finfish stocks: north-western Australia The continental shelf of north-western Australia supports a diverse and productive demersal fish community which has been exploited commercially since 1959 (Sainsbury 1987, 1988; Sainsbury, Campbell & Whitelaw 1993). Over the period that the fishery has been operating, a marked change in the species composition of the fish catch has been observed, although the total catch rate of the fishery has remained relatively constant. During the early years of the fishery, the genera Lethrinus (emperor) and Lutjanus (snapper) accounted for 40±60% of the catch, while two other genera, Nemipterus (thread-fin bream) and Saurida (grinner), together comprised about 10% (Sainsbury 1987, 1988; Sainsbury et al. 1993). However, this structure had reversed by the mid-1980s, with Lethrinus and Lutjanus comprising about 10% of the catch, and Nemipterus and Saurida around 25%. ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 406 S. J. TURNER ET AL. One explanation for the observed changes in fish community structure is that it reflects a change in habitat structure as a result of fishing activity on the shelf (Sainsbury 1987; Sainsbury et al. 1993). There has been a significant reduction in the quantity of epibenthic fauna (primarily sponges, alcyonarians and gorgonians) caught as by-catch in trawls, compared with that recorded prior to and during the early development of the fishery. The catch rate of sponges has decreased markedly (from around 500 kg h±1, but up to 2600 kg h±1, in 1963, to < 300 kg h±1, and frequently, 0 kg h±1, in 1979; Sainsbury 1987). Sainsbury et al. (1993) found that, where the fate of epibenthic organisms after impact with the trawl gear was known, epibenthic organisms were damaged (i.e. detached from the sea-bed) in 90% of observations. These changes in habitat structure were reflected in fish community structure, with Nemipterus and Saurida being recorded predominantly in open sand habitats with low densities of epibenthic fauna, while Lethrinus and Lutjanus were most common in areas with dense epibenthic fauna (Sainsbury 1987; Sainsbury et al. 1993). Campbell (cited in Coutin 1993) suggested that the direct impact of fishing mortality on the composition of fish communities is less than the indirect impact of habitat degradation or loss. Management action An `actively adaptive' approach was adopted for the management of the North-West Shelf fishery, including management experiments undertaken on a spatial scale of relevance to the fishery (Sainsbury 1987, 1988, 1991; Sainsbury et al. 1993). One of the management objectives was to assess the feasibility of maximizing the more valuable catch of Lethrinus and Lutjanus by allowing the recovery of the preferred habitat within an area of the North-West Shelf closed to commercial trawling for a 5-year management experiment. This resulted in an increase in the abundance of epibenthic fauna, as well as increases in the combined populations of Lethrinus and Lutjanus in the area closed to commercial trawling (Sainsbury et al. 1993). However, the recovery of epibenthic habitat structure after trawling ceased was slow and full recovery of the habitat following heavy trawling is predicted to take up to 20 years (Campbell, cited in Coutin 1993; Sainsbury et al. 1993). Conversely, in an area where commercial trawling continued throughout the experimental period, there was a decrease in epibenthic fauna and a decline in the catch rates of Lethrinus and Lutjanus (Sainsbury et al. 1993). 3. Deep-sea seamounts and orange roughy stocks: Chatham Rise, New Zealand With the continuing depletion of inshore fish-stocks, improvements in fishing technology (e.g. the development of fishing techniques to enable trawling of deep-sea seamounts) and the development of a new international regime for the oceans under which many coastal states have claimed extensive Exclusive Economic Zones (United Nations Convention on the Law of the Sea 1982), deep-water fishing grounds are being increasingly exploited (Probert 1996; Probert, McKnight & Grove 1997). One of New Zealand's major deep-water fishing areas is the Chatham Rise, a relatively broad submarine ridge extending for some 1000 km off central New Zealand to the east of the Chatham Islands, and rising to within 200±400 m of the surface from depths of more than 2000 m (e.g. Probert 1996; Probert et al. 1997). These deep-water ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 407 areas support abundant populations of orange roughy, Hoplostethus atlanticus Collett, hoki, Macruronus novaezelandiae Hector, and other important deep-water species, and have been fished commercially since 1978. Seamounts in the Chatham Rise region support benthic communities dominated by attached epifauna, in which large sessile species, in particular the scleractinian Goniocorella dumosa (Alcock), the stylasterid Errina chathamensis Cairns and an antipatharian, ? Bathyplates platycaulus Totton, form a distinctive and major component (Probert & McKnight 1993; Probert 1996; Probert et al. 1997). These deep-water coral communities support many associated species. While there is no quantitative evidence available, commercial fishermen noted a marked decline in the abundance, biomass and diversity of hard corals, soft corals and sponges, as well as other invertebrates, brought up in the trawls as fishing progressed in these deep-water grounds (Jones 1992; Probert 1996). Available evidence suggests that, once destroyed, these deep-water coral formations may require 200±400 years to recover (Ministry for the Environment 1997). By providing habitat structure, these deep-water communities may play a key role in the life-cycle of commercially important fish, potentially representing important areas for aggregation, courtship and/or mating, spawning grounds, or nurseries for juvenile fish. While it is unclear whether significant habitat±fisheries interactions exist in such communities, damage to deep-water sessile epifauna could have long-term impacts on the sustainability of commercial fisheries. Management action Because of the damage that mobile fishing gear is capable of causing to these deep-sea habitats, and given the likelihood that these habitat structures are potentially important in the life-history of commercial fish species, it would be prudent to adopt a precautionary approach to the management of these deep-water fisheries and protect at least some of these areas. To date, the principle legislation responsible for providing for the protection and preservation of the marine environment of New Zealand's Exclusive Economic Zone has not been used to protect any deep-water habitats from the impacts of fishing. Following a study of the seamount fauna off southern Tasmania and an assessment of the impact of deep-sea trawling, the Australian Government recently put forward a proposal to include the seamounts in what will be Australia's first deep-sea marine protected area (CSIRO 1998; Hill 1998). This follows a voluntary interim closure in mid±1995 of a 370-km2 region of the seamounts 100 km south of Tasmania to fishing and other activities which could disturb the communities associated with the seamounts. This area includes 15 out of the 70 seamounts in the region. Fisheries management and mitigation of habitat degradation or loss Fisheries management world-wide has focused (and largely continues to do so) on overfishing, with management responses directed towards restricting catch or reducing fishing effort (e.g. imposition of catch quotas, bag limits, minimum size regulations, limited entry, temporary area closures and closed fishing seasons). Such management tools were developed mainly for single-species fisheries, and little effort has traditionally been directed towards maintaining the ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 408 S. J. TURNER ET AL. ecological integrity and function of habitats and ecosystems. Fisheries management can no longer attempt simply to maximize yield, while ignoring the impacts of fishing on habitat structure and heterogeneity (Cooke & Earle 1993; Bohnsack & Ault 1996; Roberts 1997). Even low fishing mortalities will have a profound effect on habitat when caused by gear such as bottom trawls (Pauly 1997). The conservation, protection and enhancement of benthic habitat structure is increasingly regarded by the scientific community as fundamental to any attempt to rationally and sustainably manage fisheries resources in the long term (Mager & Thayer 1986; Dayton et al. 1995; Bohnsack & Ault 1996; Langton et al. 1996). The need to move from a highly focused consideration of single-species or single-habitat management strategies to integrated multispecies and ecosystem management perspectives is central to this change in management approach (Morgan 1987; Sherman 1991; Hoss & Thayer 1993; Roberts & Polunin 1993; Agardy 1994; Wilcove & Blair 1995; Roberts 1997). Habitats do not exist in isolation, nor is any one habitat used exclusively during the life-cycle of any single organism. Therefore, habitats cannot effectively be managed as separate, discrete entities. If an ecosystem management approach is to be effective, it will require a better understanding of the functioning, dynamics and interrelationships between organisms and habitats, and will need to include an appreciation of the various spatial and temporal scales over which linkages between species and habitats occur (e.g. Langton et al. 1995, 1996; Auster et al. 1997). Fisheries management needs to address habitats on a landscape scale and consider issues such as: What is the temporal and spatial sequence of habitats which are used by different life-history stages? Is there an optimum mix (i.e. combination and distribution) of habitat structure to produce optimum growth and survival of species? How does the form and extent of habitat structure influence the use by and availability to different life-history stages and species? What are the ecological relationships between the quality and quantity of habitat structure, and both fisheries production and the abundance of non-target species? (Peters & Cross 1992; Hoss & Thayer 1993). Given the numerous species, their various life-history stages, the diversity of habitats, the multitude of factors influencing the utilization of habitat structure by any one species and the complex ecological interactions among species and habitats, there is an urgent need to conserve a wide range of habitat structures and to move away from targeting single habitats for single species (Peters & Cross 1992; Auster et al. 1997). With ever-increasing demands on fish-stocks and the need for sustainable use of resources, new approaches to fisheries management are needed. Management tools for mitigating degradation or loss of habitat structure while maintaining healthy sustainable fisheries have gained widespread interest from fisheries scientists and managers. These include: (1) protective habitat management; and (2) habitat restoration. 1. Protective habitat management Protective habitat management involves the designation of `protected' marine and coastal areas (variously referred to as `marine reserves', `marine parks', `marine and coastal protected areas', `sanctuaries', `replenishment reserves' and `harvest refugia'). Protected marine and coastal areas have become a highly advocated form of marine management, providing types of ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 409 protection not offered by other management strategies (Roberts & Polunin 1993; Agardy 1994; Bohnsack & Ault 1996; Eichbaum, Crosby, Agardy & Laskin 1996; Allison, Lubchenco & Carr 1998). These areas are generally afforded some level of protection from fishing (ranging from being totally non-extractive areas to areas of restricted harvesting), and have as their ultimate goals (among other considerations) the sustainable management and use of fisheries resources, the conservation of benthic habitat structure and biodiversity, and the safeguarding of ecosystem function and integrity (Jones 1994; Ticco 1995; Bohnsack & Ault 1996). Protected areas are especially important in providing protection for habitats which are critical to the viability of a species or population, such as spawning grounds, nursery grounds and those which are centres of high species diversity. They range in size from small, closed areas or harvest refugia, designated to protect a specific resource or habitat type, to extensive areas which are designed to integrate the management of many species, habitats and uses in a single comprehensive ecosystem-based management plan (Agardy 1994; Eichbaum et al. 1996). As fisheries management tools, protected marine and coastal areas are primarily intended to enhance or replenish fisheries yields through a reservoir of broodstock. These areas also provide for preservation of habitat diversity and integrity by protecting habitat structures from damage, providing sources of recruits or sinks for enhanced settlement of those species important in contributing to habitat structure, preserving genetic diversity within species, and maintaining more natural population and community structures (cf. Carr & Reed 1993; Dugan & Davis 1993; Roberts & Polunin 1993; Bohnsack & Ault 1996; Auster et al. 1997). Work on the North-West Shelf fishery of Australia has demonstrated the effectiveness of large area closures to fishing, and has shown that the impacts of fishing on epibenthic habitats and associated fish assemblages do not occur in areas where commercial fishing is stopped (Sainsbury 1987, 1988, 1991; Sainsbury et al. 1993). Central to arguments advocating the development of protective habitat management is the idea that, unlike traditional fisheries management strategies which require large amounts of information about the life-histories of the species being fished, their implementation should not be dependent on the acquisition of large amounts of information about the life-histories of each species, and their interactions with habitat structures and other species (e.g. Roberts & Polunin 1993; Auster et al. 1997; Allison et al. 1998). Rather, by protecting some habitat structure until a full understanding of the effects of fishing impacts can be determined, the risks of total loss of habitat and of associated resources are minimized. Furthermore, areas selected for protection need not be permanent or immutable, but can be adjusted as data improve (Roberts 1997). Management for habitat structure and heterogeneity by establishing areas closed to mobile fishing gear is consistent with the precautionary approach and `risk aversive' management strategies (e.g. Cooke & Earle 1993; Agardy 1994; Jones 1994; Dayton et al. 1995; Bohnsack & Ault 1996; Eichbaum et al. 1996; FAO 1996; Auster et al. 1997; Roberts 1997). Additionally, protective habitat management strategies provide a buffer against scientific uncertainty and/or difficulties in executing management measures (e.g. Dugan & Davis 1993; Bohnsack & Ault 1996; Roberts 1997; Allison et al. 1998; Lauck, Clark, Mangel & Munro 1998). Fundamental research still needs to be undertaken to determine the ideal location of protected areas, as well as the size, number and total area required to achieve specific ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 410 S. J. TURNER ET AL. management goals (Tisdell & Broadus 1989; Salm & Price 1995; Bohnsack & Ault 1996). This will require integrating knowledge from many different disciplines. In addition, the proportion of habitat which should be protected is an issue that needs to be resolved. The US South Atlantic Fishery Management Council recommended that 20% of the total shelf area should be set aside as marine fisheries reserves, based on the goal of maintaining stock reproductive output at levels of 30% or more of that of an unexploited population (Bohnsack 1990, cited in Roberts 1997). Ballantine (1991) recommended that 10% of all marine habitats in New Zealand be designated as non-extractive marine reserves. While Cooke & Earle (1993) suggested: `Whether or not there is evidence of adverse effects, any fishing method used in an area that involves substantial disturbance to habitat shall be excluded from representative closed sub-areas covering at least 50% of the fishing ground, to conserve part of the habitat in its undisturbed state.' Furthermore, the above authors suggested that, if the entire fishing ground had been subject to major disturbance, closed areas should be established to allow habitat recovery. Sladek Nowlis & Roberts (cited in Roberts 1997) considered that larger protected areas (up to 50% or more) may continue to provide additional benefits, especially in heavily exploited fisheries. Even the most well-designed and managed protected marine and coastal areas cannot alone protect habitats, and thus, fisheries against anthropogenic impacts outside their boundaries (ranging from local pollution events to widespread or global phenomena such as persistent widespread pollutants, episodic climatic events, disease epidemics and the spread of exotic species). Without adequate protection of species, habitats and ecosystems outside the boundaries of protected areas, the effectiveness of protected marine and coastal areas will potentially be severely compromised (Carr & Reed 1993; Agardy 1994; Jones 1994; Allison et al. 1998). Protected areas should not be seen as a solution replacing conventional management approaches, but need to be one component of an integrated programme of coastal zone and fisheries management (Roberts 1997; Done & Reichelt 1998). 2. Habitat restoration Habitat restoration seeks to re-establish important habitat structures and their ecological function to a level, based on feasibility and historical information, which ensures the greatest long-term continued productivity of the system. Habitat restoration is usually embarked upon after all the alternatives to prevent or mitigate adverse impacts have been unsuccessful. While the restoration of natural communities is an established method in land and freshwater management, restoration of marine systems has only recently received attention. In the past, the focus of marine ecosystem restoration has been the curtailment of the source(s) of degradation or loss, and typically, the impacted communities have been left to recover naturally. Nevertheless, natural recovery of impacted areas can be very slow since many important habitat-forming species are slow colonizers, and may take a long time to attain natural population densities, size and age structures. Ecosystems often do not recover from anthropogenic disturbance without additional management or manipulation (Pratt 1994). Where human-induced impacts have resulted in the loss of habitat structure, associated changes in the hydrodynamic, physical and/or chemical conditions make habitat restoration ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 411 more difficult, and in the majority of cases, the original environmental conditions will need to be re-established before habitat restoration can be successfully achieved (e.g. Thayer, Fonseca & Kenworthy 1986; Fonseca, Thayer & Kenworthy 1987; Kirkman 1992). Proper site selection, species selection and the timing of restoration effort are of paramount importance to the success of restoration projects (e.g. Fonseca et al. 1987; Fonseca 1989). Habitat restoration should not only focus on the physical habitat type, but also its ecological functions, and the success of any restoration project should be measured not only by the persistence of the habitat, but also the extent to which the habitat functions and is used by organisms which would normally be found there (e.g. Fonseca et al. 1988; Fonseca 1989; Thayer, Fonseca, Kenworthy, Colby & Currin 1989; Thayer, Fonseca & Kenworthy 1990; Levings 1991; Pratt 1994). Ignoring ecological linkages may result in a poorly functioning ecosystem with low probability of persistence (Pratt 1994; Race & Fonseca 1996). As with the establishment of protected marine and coastal areas, the restoration of habitat structure must be addressed in the context of integrated coastal zone management policies if the goal of habitat restoration is to be achieved. Protected marine and coastal areas may also exist as restoration areas, allowing ecosystems to return to a condition more closely approximating the natural state in terms of habitat quality, species abundance and diversity, population structure, and reproductive output (Irving 1994; Lindeboom 1995). In the future, the establishment of protected areas selected for their potential to be restored could usefully complement networks of areas set up to conserve habitats (Gubbay 1995; Done & Reichelt 1998). Although techniques exist to restore salt marshes, seagrass beds, mangroves, kelp forests, coral reefs and temperate reefs (see Thayer 1992), in many cases, the process of restoration has not been overly successful nor have there been any long-term evaluations of restoration success (Pease, Thayer & Lillestolen 1994). Furthermore, there is currently little evidence that restored habitats are as capable of supporting or sustaining the ecological function and fisheries production which their natural counterparts do. While studies clearly show, for example, that increasing habitat structure and heterogeneity using artificial reefs can increase the local abundance of fish, it has not been conclusively demonstrated that the total fish yield for any body of water increases significantly as a result of structural modification, rather it may be that the artificial habitat tends to concentrate fish that were already present (Bohnsack 1989, 1991; Ryder & Kerr 1989; Sheehy & Vik 1992). There is also little evidence that restored or created seagrass and salt marsh habitats function ecologically in the same way as their natural counterparts. The time required for equivalent functioning of restored or created habitats may be on the scale of years (e.g. Homziak, Fonseca & Kenworthy 1982; Smith, Fonseca, Rivera & Rittmaster 1988; Fonseca, Kenworthy, Colby, Rittmaster & Thayer 1990; Meyer, Fonseca, Colby, Kenworthy & Thayer 1993). Meyer et al. (1993) reported that these habitats were still not fully functional in terms of living resource utilization even after 2 years of created marsh and 3 years of created seagrass development,. Because of a fundamental lack of understanding of the ecology and dynamics of natural habitats and their biological resources, as well as lack of detailed understanding of the biology of individual species in restored ecosystems, restoration of marine systems has not yet evolved into a reliable management tool. The science of habitat restoration is still `young, imperfect ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 412 S. J. TURNER ET AL. and experimental', and in most cases, the extent to which restoration compensates for the loss of natural habitat structure, the trade-offs that occur if out-of-kind restoration is undertaken or the rate of functional replacement if restoration is successful, are unknown (Thayer, Fonseca & Kenworthy 1985; Thayer et al. 1990; Race & Fonseca 1996). Since habitat loss still cannot be controlled, Fonseca et al. (1988) advocated that, at least in the case of seagrass beds, complete conservation of seagrass habitat should be undertaken as the alternative to restoration. Fisheries law and policy A number of recent international fisheries agreements have recognized the effects of fishing on habitat as a key management issue, and specifically identify the need to provide for habitat protection and restoration to ensure the long-term sustainability of fisheries resources (Garcia & Newton 1994; Hey 1996). These instruments include the 1995 International Code of Conduct for Responsible Fisheries (FAO 1995) which sets out principles and standards for responsible fishing to ensure effective conservation, management and development of all living aquatic resources with due consideration for the ecosystem and biodiversity. The Code includes a specific requirement that `critical fisheries habitats' are protected from destruction, degradation, pollution and other significant impacts resulting from human activities which threaten the health and viability of the fishery resources (Article 6.8). In addition, the Code provides that every effort should be made to ensure that fisheries resources and habitats critical to the well-being of such resources which have been adversely affected by fishing or other human activities are restored (Article 7.6.10). The Code also seeks to improve fishing practices. Of particular relevance to the protection of habitat, is the requirement to ensure that assessments of the implications of habitat disturbance are carried out prior to the introduction to an area of new fishing gear, methods and operations on a commercial scale (Article 8.4.7). New approaches to fisheries management, including the development of protected marine and coastal areas, and habitat restoration, are also increasingly common components of fisheries management programmes under fisheries law and policy at a national level. The Magnuson±Stevens Fishery Conservation and Management Act is the United States' foremost federal law governing fishing activity in the 200-nautical mile Exclusive Economic Zone. The 1996 amendment to the Act (the `Sustainable Fisheries Act') addresses a number of major fisheries management issues, including that of habitat degradation and how to strengthen provisions for habitat protection and restoration (Magnuson-Stevens Fishery Conservation and Management Act 1996). The Act recognizes that some fish stocks have declined to the point where their survival could become threatened as a consequence of direct and indirect habitat losses which have resulted in a diminished capacity to support existing fishing levels [Section 2(a)(2)], and that one of the greatest long-term threats to the viability of commercial and recreational fisheries is the continuing loss of habitat [Section 2(a)(9)]. The new Act mandates guidelines be established for the description and identification of `essential fish habitat', as well as the adverse effects of activities on such habitat (including impacts from fishing), and in the consideration of actions to ensure the conservation and enhancement of such habitat ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 DEGRADATION OR LOSS OF HABITAT STRUCTURE 413 [Section 305(b)(1)(A)]. Fishery management plans are required to include management measures which minimize the adverse effects on habitat caused by fishing and must include an assessment of the potential adverse effects of all fishing gear types used where there is essential fish habitat [Section 303(a)(1)]. The Act also provides for fisheries research to be undertaken to develop and test new gear technology and fishing techniques to minimize any adverse effects on essential fish habitat [Section 404(c)(2)]. Similar developments in fisheries management have been seen elsewhere. The new New Zealand Fisheries Act was enacted in 1996 to provide for the utilization of fisheries resources while ensuring their sustainability, and at the same time, avoiding, remedying or mitigating any adverse effects of fishing on the environment (Section 8) (New Zealand Fisheries Act 1996). Under the Act, fisheries management decisions must take into account a number of `environmental principles', including maintenance of the long-term viability of species which are associated with or dependent upon the harvested species, maintenance of biological diversity, and protection of habitats of particular significance to fisheries management (Section 9). The Act provides for the setting or varying of `sustainability measures' for stocks or areas, after taking into account any effects of fishing on the environment, among other factors [Section 11(1)(a)]. Thus, the Act sets out to ensure that account is taken of the wider effects of fishing in the ecosystem in managing fisheries resources. Likewise, the Queensland Fisheries Act provides for the management, use, development and protection of fisheries resources and `fish habitats' within the coastal waters of Queensland, Australia, with the objective of ensuring that fisheries resources are used in an ecologically sustainable way [Section 3(1)(a)] (Queensland Fisheries Act 1994). The Act provides for the protection and conservation of fish habitats through the declaration of `fish habitat areas' which are managed under management plans (Sections 120±122). There are also provisions for the rehabilitation or restoration of fish habitat (Sections 124±125). The objectives of the Western Australian Fish Resources Management Act similarly include the conservation of fish and protection of their environment, and the designation and management of `fish habitat protection areas' [Section 3(2) and Sections 115±119] (Western Australian Fish Resources Management Act 1994). Regulations relating to the protection or management of fish habitat protection areas may prohibit or regulate fishing or any other activity which may adversely affect the designated area [Section 120(2)]. While there is undoubtedly a consensus on the substantive issues to be addressed in fisheries management, as reflected in international agreements and recent developments in fisheries law and policy at a national level, the question remains whether sufficient political determination, funding and resources exist to do justice to these important and ambitious undertakings to prevent further habitat degradation or loss. Conclusions As more fisheries species are exploited and technological advances increase the range of habitats which are likely to come under threat from fishing, understanding and effectively managing for the effects of fishing on habitat structure becomes ever more important. There is a growing international consensus that a `precautionary approach' should be adopted towards fisheries management, and that this includes a duty to assess the impact of fishing on the ã 1999 Blackwell Science Ltd, Fisheries Management and Ecology 1999, 6, 401±420 414 S. J. TURNER ET AL. environment, and to adopt management strategies to protect and restore habitats of special concern. 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