Goose ( per km2) and habitat area (km2) estimated using a GIS. These estimates are ( polymerus) attach then adjusted using the CPUE for each area. themselves by flexible stalks in dense Two area-swept trawl surveys have been clusters on mid rocky intertidal shores completed; central west coast of Vancouver exposed to high wave action. Life history of Island (Nootka region) in 1999 and Queen goose in British Columbia has Charlotte Sound and West Coast of Haida been recently reviewed by Lauzier (1999 a). Gwaii in 2000. Approximately 40% of the coast has been covered by the industry trap Landings for goose barnacle are recorded survey. At present, DFO is working with once from PFMA 1 (1994) and three times in industry to complete the trap survey. PFMA 2 (2E + 2W) between 1988 and 1993. Where the trap survey has been completed, Total landings from all areas in all years DFO is initiating an experimental fishery to were <10 t. As with king crab, these were assess population responses to fishing and exploratory fisheries and there has been no validate biomass estimates (Workman et al. further fishing since 1994. The fishery is 2000). The current experimental fishery is presently closed in British Columbia. operating off the northwest coast of Vancouver Island in PFMA 125. Tanner Crabs In summer 2000, exploratory sounding, Tanner ( tanneri and C. trawling and trapping was undertaken in angulatus) are deep-water continental shelf the deep continental slope waters off Queen and slope predatory species. There are Charlotte Sound and the West Coast of established fisheries for Chionoecetes species Haida Gwaii. Survey objectives were to in Alaska and in eastern Canada where they identify potential Tanner crab habitat, are called “snow” crab. The life history of conduct experimental trawling and Tanner crab is little known from British trapping for distribution and abundance Columbia waters. data and collect general species biodiversity data from these unfished depths. Fisheries and Oceans Canada began Commercial fishing has taken place in these investigating the commercial potential of areas previously, but at much shallower Tanner crab (Chionoecetes tanneri and C. depths. Trawling and trapping (at 500 to angulatus) with a literature review and 1900 m depth – Figure 31) was executed development of an exploratory assessment with a large trawl (fitted with a program in 1997. Assessment and small-mesh cod-end liner to retain development of a possible future to 10 mm length) or with baited conical top- commercial fishery on these species is loading traps. In traps, crabs dominated the occurring under the science advisory catch with fewer than a dozen species framework (Perry et al. 1999) for new and encountered as by-catch, conversely, in developing invertebrate fisheries (J. trawls >140 taxa were captured in Boutillier and G. Workman, DFO, personal association with crabs. Each set’s catch was communications). Two sources of data are sorted to species and weighed, enabling the required to assess Tanner crab; area-swept identification of all animals. For all species trawl density data collected by DFO and of crab, size, weight, maturity state, and distributional trap Catch Per Unit Effort shell condition were recorded for use in (CPUE) data collected by industry stock assessments and in life history studies. participants. Biomass estimates are Deep sampling yielded increased catches of calculated using the trawl density data C. angulatus and two other little known deep water crab species, Paralomus verrilli

110 111 and P. multispina. Sufficient numbers of P. Sea Cucumber verrilli and P. multispina were caught to provide the first basic population data ever The red sea cucumber (Parastichopus collected for these species in British californicus) is a large species (to 45 cm Columbia. At depths >1500m P. verilli length) found from the lowest intertidal dominated, from 1000 to 1500 m depth C. zone to over 100 m depth on a wide range angulatus and P. multispina dominated, and of rocky to sandy habitats of moderate to at depths <1000 m C. tanneri and P. low exposure. It uses a ring of cauliflower- multispina were most abundant. Tanner like tentacles at the anterior end of its crab (C. tanneri) were less abundant off cucumber-shaped body to mop up Haida Gwaii than off the West Coast of deposited material. Sea cucumbers digest Vancouver Island. Significant quantities of the organic components of ingested C. angulatus, however, occupy the deeper sediments and void the indigestible depths off Haida Gwaii. The preliminary components. The life history of this species invertebrate species biodiversity data (»70 is poorly known (Boutillier et al. 1998). taxa) were contributed to the database in Appendix D and the full cruise results will Commercial fishers dive to collect sea be reported by Workman et al. (2000, in cucumbers by hand. Catch weight is preparation). reported “split” (de-watered and gutted). Products are boiled body walls for export to Green Sea Urchin Asia and muscle strips from inside the body walls for both export and domestic Green sea urchin (Strongylocentrotus consumption. The management of this droebachiensis) is a circumpolar, arctic-boreal fishery is covered in detail the Integrated species whose life history is particularly Fishery Management Plan, so only a Haida well studied in the north Atlantic. It is a Gwaii area overview is provided. smaller species than the red sea urchin, but is also a herbivorous grazer on attached and The first landings of sea cucumber from drift seaweed. Fishing is by divers and the Haida Gwaii started in 1987 from PFMA 2E commercial product is sea urchin roe. As and 2W. From 1987 through 1995, PFMA with red sea urchin, sea otters are known to 2W reported total landings of three t split reduce green sea urchin populations in weight from two years of fishing whereas Alaska and British Columbia (Watson and PFMA 2E averaged 42.5 t annually from Smith 1996). nine years of fishing. Since 1996, Haida Gwaii has been closed to fishing due, in The fishery in Haida Gwaii has been small part, to low market value and processing with exploratory landings from PFMA 1 in logistics rendering this area economically 1989 and 1990 and PFMA 2E in 1990 only. marginal. The other reason is that sea Combined north coast (PFMAs 1 to 10) cucumber is the target of the first shellfish landings between 1987 to 2000 have not adaptive management experiment in British exceeded 150 t. Coast-wide, the fishery is Columbia as described in the Integrated dominated by south coast landings (Perry Fishery Management Plan. Adaptive and Waddell 1999). All of Masset Inlet management is discussed below. The Haida (PFMA 1-6) is closed to commercial fishing Gwaii closure is part of such an experiment, as an allocation to Haida food fishing now halfway through its 10-year life. It is (Figure 29). possible that the fishery could be reopened for Haida Gwaii at some time after this experiment.

112 SHELLFISHERIES MANAGEMENT • each fishery is unique in the blend of SUMMARY biological and management criteria used; Shellfisheries management is a moving • DFO attempts to provide a scientific target involving multiple species with basis for all management advice; and criteria changing according to fluctuating • not all fisheries depend on stock stock status and new technical information. assessments for their management. Nonetheless, an overview of the basics is provided here. The major shellfisheries The species’ life histories and the human occur mostly inshore with the exception of understanding thereof dictate the in the northwestern region management criteria and will influence of Hecate Strait (Table 24). Among those future spatial management decisions. For with offshore landings (Dungeness crab and Gwaii Haanas, management of the most red sea urchin), almost 100% of landings are important fisheries is based on total in the east and northern offshore (PFMAs allowable catches arising from stock 101 and 102) compared to the west and assessment (geoduck and red sea urchin) southern offshore (PFMAs 130 and 142). and the spawner index. There is a high local (HFP) labour input into stock With so much shellfisheries information, a assessments for the diving fisheries. quick management overview is warranted. Summarized in Table 25 are the Other issues relevant to fishing warrant management criteria for the major mention as follows: shellfisheries of Haida Gwaii. The main points are as follows: • Land-based pollution - the low population levels of Haida Gwaii region with attendant low levels of land-based

113 pollution render habitat degradation • Sea-based pollution – this is a genuine and pollution minor issues compared to threat, such as oil pollution from a some urban and industrial mill- shipping incident offshore or from an oil associated south coast areas of British or gas production platform blowout Columbia (Orensanz and Jamieson should development of the known 1998). For example, there has been no reserves of Hecate Strait occur. Spilled logging in the Gwaii Haanas area since oil can have significant short-term acute 1987, no other industrial upland activity affects, long-term chronic effects and oil for decades and there will always be can persist for decades in sheltered very little built infrastructure in the area. marine shore habitats (Sloan 1999). Elsewhere in Haida Gwaii, logging is Recoverable resources are estimated at active and there are downstream effects »414 million m3 [2.6 billion barrels] of oil such as increased sedimentation in and »565 billion m3 [20 trillion ft3] of gas nearshore waters. Coal and mineral in Hecate Strait (Dietrich 1995). There exploitation were important post- have been federal and provincial contact exploration and settler moratoria on exploration in waters attractants (Richardson 1873; Morton around Haida Gwaii since 1972. In 1992). There has been no mining in the 2001, a new British Columbia Gwaii Haanas area since 1968 when the government wanted to address lifting Jedway Iron Mine (Harriet Harbour) the provincial moratorium in favor of closed. The largest nearby regional regional economic development mine was in Tasu Sound and it closed in (Observer, July 2001). Debate on the oil 1983. and gas issue is about to intensify. Most of the Gwaii Haanas area itself has low petroleum resource potential (Dietrich et

114 al. 1992). Further, the West Coast bioherms and coral and sea fan groves. Offshore Exploration Environmental These impacts are not quantified but Assessment Panel recommended that there is genuine concern for deep-water possible future drilling should be corals throughout temperate regions prohibited within an exclusion zone of world-wide (Risk et al. 1998). The whole 20 km from any point of land issue of conservation of deep-water (WCOEEAP 1986, p. 93). The landward invertebrates is grossly underdeveloped. edge of this zone, therefore, would be seaward of the proposed Gwaii Haanas • New invertebrate fisheries – some marine boundary [established in the fisheries, such as geoduck, pre-date the South Moresby Agreement, 1988] by »10 intention to create the Gwaii Haanas km. Finally, Gwaii Haanas is further national marine conservation area as protected in that jurisdiction over declared in the South Moresby Agreement seabed lands within the proposed Gwaii (1988) between the province of British Haanas marine area was transferred Columbia and Canada. Other fisheries, from British Columbia to Parks Canada however, were initiated in the Gwaii by provincial Order in Council in April Haanas area after that, such as red sea 2001. All other types of water-borne urchin. Still other fisheries are under pollution are less threatening to Haida consideration for development such as Gwaii than to, for example, the densely Tanner crab. Once the Gwaii Haanas populated and enclosed southern Strait marine area is declared, prospects for of Georgia. new fisheries will be important management issues into which the • Habitat-modifying fisheries – these regional Gwaii Haanas management include heavy bottom trawls used by partnership will contribute. “draggers” in demersal (near-bottom) finfisheries. Watling and Norse (1998) • Access to full fisheries information – the estimated the amount of sea bottom long-term sustainable, ecosystem-based scoured by trawls world-wide annually management envisioned for the at an astonishing half of Earth’s eventual Gwaii Haanas management continental shelf or the equivalent of 150 partnership will require access to ALL times all the forest land area clearcut possible data sources on marine annually! The extent of damage to resources. At this time, for example, invertebrate benthic communities by Parks Canada has no access to the trawls in the Haida Gwaii region is confidential fisheries data such as unknown. Heavy trawl gear is not used geoduck fishing areas. Mechanisms will in regional shellfisheries in Haida have to be developed by which Gwaii Gwaii. Some habitat modification does, Haanas management will have access to however, take place in the sediment such data while maintaining disturbance by geoduck divers and in confidentiality. There is precedent as shrimp trawling with light-weight Gwaii Haanas retains confidential data trawls. Their scale and intensity does on Peregrine falcon (Falco peregrinus) not compare with the finfish draggers, eyries (nests) through an arrangement and, to a lesser extent, finfish long- with the province of British Columbia. liners. In the Haida Gwaii region, it is This information can be made available likely that finfish dragging and long- from Gwaii Haanas if written lining does impact deep-water permission is first received by Gwaii invertebrate populations such as sponge Haanas management from the province.

115 SOCIOECONOMIC OVERVIEW OF figures are listed in Table 26 for the main HAIDA GWAII SHELLFISHERIES shellfisheries of Haida Gwaii. Although only landed every three years, geoduck is “Systematic social studies should be conducted by far the most valuable shellfishery. Gwaii to accurately evaluate the impacts of a proposed Haanas area landed value dominated in the MPA on community stability.” (NRC 2001) geoduck and the prawn trap fishery. The domination by dive fisheries of shellfish Science and management of these landed value is coast-wide (Harbo 1998). In shellfisheries are but a part of the picture. 2000, a geoduck fishing year, total shellfish Also needed is a detailed socioeconomic landings from Gwaii Haanas alone assessment of their impacts to Haida and exceeded $14 million. This is an important other communities in Haida Gwaii, the number to keep in mind. In public north mainland coast and the whole south consultation concerning DFO’s Gabriola coast of British Columbia. The fishers, Pass marine protected area, a win-lose packers, processors and support services all polarity developed between the geoduck make appreciable contributions to their industry and an environmental NGO home communities. Important questions (Spisak 1997). It will be essential for the need to be answered on who benefits from Gwaii Haanas consultation process to be these shellfisheries and who could be proactive in ensuring that all sectors negatively impacted by the establishment of contribute, enabling a civil and equitable the Gwaii Haanas marine conservation area. balance of sustainable resource extraction with appropriate conservation area A simple way to view the gross economic management. value of shellfisheries is to look at their annual landed value. The most recent The issue of where the licence holders live relates importantly to economic benefits

116 from shellfisheries. Listed in Table 27 are south coast). These represent a process the home locations of commercial licence- under the Aboriginal Fisheries Strategy to holders according to various Haida Gwaii allocate more licences regionally to First shellfisheries. A reasonable assumption is Nations. that most ships’ crews tend also to come from the licencees’ home areas. With the Shellfish processing is an important shore- exception of razor clam and Dungeness crab based economic attribute of shellfisheries fishers, few commercial licence-holders live contributing employment and development in Haida Gwaii. Such information is in coastal communities. Most landed important as income from fishers residing shellfish passes through licensed processing elsewhere is less likely to contribute to plants. for export (e.g., all geoduck, communities near where they fish. Added all sea urchin, most ) must pass to that could be the importance of through a “federally registered” plant to employment which is currently depressed acquire an “export certificate.” These plants in coastal communities. An important trend are inspected regularly by the Canadian is that of “communal” (F) licences. These are Food Inspection Agency (CFIA) for licences converted from the general pool compliance with seafood safety regulations and reallocated to First Nations. There are (CFIA 1999). Seafood for domestic now five communal red sea urchin (FZC) consumption only (e.g., some prawns and licenses (four north mainland coast / one Dungeness crab) passes through Haida Gwaii), three communal crab (FR) provincially-registered plants inspected by licences (two Haida Gwaii / one south the British Columbia Ministry of Fisheries. coast) and two communal prawn (FW) Shellfish can be landed directly to licences (one north mainland coast / one processors by fishers or delivered to

117 processors by packers in a variety of states. high value province-wide, this preliminary The Haida Gwaii prawn catch, for example, assessment reveals that, with exceptions for is frozen at sea whereas red sea urchin must razor clam and Dungeness crab, most reach processors fresh for specialized economic benefits from commercial fishing handling, and geoducks must go through bypass Haida Gwaii communities. processing and reach their Asian export markets live. The prawn catch destined for PUBLIC HEALTH AND SAFETY OF export (Japan) is treated on board with a SEAFOOD Japanese anti-oxidant to maintain bright colouration. This chemical is not CFIA- There are public health risks associated with approved for domestic use, so the product eating marine invertebrates in Haida Gwaii. can only be exported. The data in Table 28 Bivalve seafood safety is managed under show that, with the exception of all razor the Canadian Shellfish Sanitation Program clam and 20% of Dungeness crab with the following agency mandates: processing, product tends to be processed off Haida Gwaii. Dungeness crab had the • biotoxin surveillance and seafood most complex processing geography with inspection by the Canadian Food appreciable proportions of catch being Inspection Agency; processed in Masset and Prince Rupert. • bacteriological water quality surveys by Although the Masset plants are federally Environment Canada; and registered, they do not participate much in • posting and patrolling of closed areas the overseas export certification of shellfish. and the management of conditionally approved areas by DFO under the In summary, although the shellfisheries of Fisheries Act – usually on Haida Gwaii (and Gwaii Haanas) are of

118 recommendation from the above cramps and difficulty breathing. These is agencies. no drug treatment; care-givers should call emergency response, induce vomiting and The Canadian Shellfish Sanitation Program stand-by to initiate cardio-pulmonary arose from a Canada-U.S. Bilateral resuscitation (CPR) to assist breathing if Agreement observing equivalent standards necessary. and is routinely audited by the U.S. Food and Drug Administration (EC 1997). The Toxic red tides are well known in British Sanitation Program’s objectives are to Columbia (Quayle 1969; Martin, 1994) and minimize the public health risk posed by Alaska (RaLonde 1996). They are also consumption of Canada’s bivalve mollusc known in traditional Haida stories as seafood. Dungeness crab seafood safety described previously in the Haida use does not fall under this program, but is section. Given the frequency of red tides included in Canadian Food Inspection along the north coast of British Columbia Agency surveillance. and the lack of regular PSP testing, the whole region (PFMAs 1 to 10) has been Paralytic Shellfish Poison (PSP) Biotoxin closed by DFO to all intertidal bivalve fishing since 1963 for public health and Paralytic Shellfish Poisoning (PSP) is caused safety. Gwaii Haanas visitor management by several neurotoxins in certain policy supports this closure year-round. dinoflagellate phytoplankton species. Areas, such as North Beach on Graham Many species of dinoflagellates can form Island, are opened for fishing only after seasonal (spring/summer) aggregations testing. The internationally recognized and (“blooms”). Visible blooms are called “red approved “mouse bioassay” test, tides” because of their red-brown administered by the Canadian Food discolouration of seawater. The blooms are Inspection Agency is used. The test patchy in distribution and their timing and threshold is a toxin level of >80 mg toxin per locality are very unpredictable. 100 g of bivalve tissue. For a closed area to be reopened, the above toxin level cannot be Bivalve molluscs such as clams and mussels exceeded in any of three samples taken over consume phytoplankton by filter-feeding. a 14 day period. The geoduck industry When there is a bloom of toxic covers the costs of PSP testing of product dinoflagellates, bivalves ingest them in from Haida Gwaii before full-scale fishing large numbers. Toxins are retained in during the years in which they fishing. bivalve species for varying periods of time. Subtidal geoduck results cannot be used to People and other warm-blooded vertebrates open adjacent intertidal areas for clam can be poisoned after eating such digging. The Haida Fisheries Program contaminated shellfish. The toxin dynamics expedites regular PSP testing to maintain vary as to bivalve species, for example, the North Beach (sub-area 1-5) razor clam butter clams (Saxidomus gigantea) absorb commercial and recreational fishery toxins gradually and retain them for longer openings. periods, whereas mussels (Mytilus spp.) absorb toxins rapidly, but tend not to retain Amnesic Shellfish Poison (ASP) Biotoxin them for as long. The toxin can be fatal through paralysis of breathing muscles Amnesic Shellfish Poisoning (ASP) is a causing inability to ventilate the lungs. relatively new biotoxin threat. It was first Symptoms are tingling extremeties, described from eastern Canada in 1988 dizziness, nausea, vomiting, abdominal where toxic mussels (Mytilus sp.) poisoned

119 over 100 persons, a few of them fatally (Ahmed 1991). It is also a neurotoxin (domoic acid) absorbed by bivalves from their phytoplankton (diatom) food. Locally, the toxin has also been found in the digestive organ (hepatopancreas) of Dungeness crabs, perhaps because they are bivalve predators. Toxic symptoms include vomiting, abdominal cramps, diarrhea, disorientation and memory loss particularly of short-term memory. Amnesic Shellfish Poisoning has been recorded from north coast of British Columbia bivalves since the early 1990s, but levels have been generally low. In eastern Canada, levels of >20 mg toxin per one gram of mussel tissue have induced fishing closure. In Haida Gwaii, there has been one ASP closure (North Beach – PFMA sub-area 1-5) for 10 months in 1995-1996 due to toxin levels recorded from razor clam and Dungeness crab.

Shellfish Growing Water Quality (fecal contamination)

Fecal coliform bacteria contamination of Haida Gwaii (including Gwaii Haanas) marine waters is monitored triennially by Environment Canada’s Shellfish Water Quality Protection Program executed by the Environment Protection Branch, Pacific and Yukon Region. Fecal contamination can come from human sewage or wildlife feces. Sampling is part of the delivery of the Canadian Shellfish Sanitation Program. If coliform counts are too high, Environment Canada recommends bivalve closures for implementation by DFO under the Fisheries Act. For closed areas to be reopened, DFO relies on Environment Canada recommendations based on sample results. Near Haida Gwaii communities, coliform levels have been sufficiently high for DFO to enact the closures illustrated in Figure 32.

120 121 CONTRIBUTION OF MARINE achieving acceptance for integrating INVERTEBRATE ISSUES TO GWAII monitoring costs into long-term operational HAANAS’ MANAGEMENT funding – not short-term research funding.

MARINE MONITORING Although we are in the earliest stages of understanding Gwaii Haanas’ marine “Without reliable data about rates of change ecosystems, there is sufficient knowledge within any habitat, there is no possibility that and experience to assess our monitoring we can predict the sorts of changes that are likely needs. In the long-term, monitoring will be to be associated with human interferences and a foundation of Gwaii Haanas’ marine developments.” (Underwood and Kennelly management. And, as invertebrates 1990) comprise »90% of our marine fauna, they will be fundamental to any monitoring “Long-term ecological monitoring is the first program. An overall monitoring program step in learning how to assess ecosystem should be vetted through interagency health.” (Davis 1993) (Parks Canada, DFO, EC) and public consultation because the data would be a “We need long-term data sets if we are to shared regional marine asset. Ideally, understand this dynamic world and our impact appropriate monitoring would both assist upon it. We may not be able to make precise Gwaii Haanas management and feature predictions about future events, but Gwaii Haanas as a major marine understanding the past will shed light on our environmental reference location for Pacific ability to alter natural processes.” (Bondrup- Canada. Nielsen and Herman 1995) Monitoring should be treated as part of “…. effective scientific research and monitoring adaptive management. This is a structured programs must be developed together” (Murray process of “learning-by-doing” that treats et al. 1999) management as an experiment in which hypotheses are formulated and against “Well-designed long-term monitoring programs which findings are used to test these will be necessary to gather data about the hypotheses. Included in an adaptive pathways of population and ecosystem management regime are target variables, rebuilding, to assess benefits, to increase pre-set values and decision points knowledge of both fishers and scientists, and to (management options) established in improve the level of protection.” (Sumaila et al. advance depending on whether explicit 2000) performance criteria are met. Monitoring facilitates the feed-back necessary to guide Monitoring is a fundamental component of adjustments as the experiment unfolds and Parks Canada’s mandate for ecosystem instructs managers. Monitoring should, integrity in terrestrial national parks therefore, be experimental and begin with a (Woodley 1993; Parks Canada Agency 2000) conceptual model of the ecosystem and be and for facilitating sustainable use without focused on the population dynamics of compromising the ecosystem structure and selected species relative to key ecosystem function in marine conservation areas. One components and physio-chemical of the great benefits of protected spaces environmental variables (Davis 1993). such as national parks is their role as long- term regional benchmarks of environmental Although we cannot monitor all species, we well-being. An important challenge is should include as many as possible because

122 they all occupy different niches. Reliance of the U.S. marine monitoring commitment on a few selected species may not reveal as follows: meaningful changes in variables such as species diversity. Examples are that mussel • indicating ecosystem health; densities may remain, but the numbers of • defining limits of normal variation; species living within the mussel bed • identifying abnormal environmental infrastructure may vary with different conditions; and conditions or that important differences • verifying agents of abnormal change. between under-rock and on-rock species may occur while one of these communities Channel Islands National Park, CA is the remains relatively unchanged. U.S. National Park Service marine inventory and monitoring program model for non- In a major marine area conservation review destructive, fixed-site monitoring (Davis et of science (NRC 2001), the following three al. 1994; Davis et al. 1997; Dye 1997 – U.S. tasks were considered central to monitoring: National Park Service inventory and monitoring program: http:// • assessing management effectiveness; www.aqd.nps.gov/natnet). Also • measuring long-term trends in committed to monitoring on the Pacific ecosystem properties; and coast is the U.S. National Marine Sanctuary • evaluating economic impacts, program (Monterey Bay National Marine community attitudes, involvement and Sanctuary 1999, 2000). California remains compliance. the leading Pacific state for marine monitoring (Murray et al. in press). To these we would add maintaining Examples are provided below using some of scientific accuracy and repeatability. Channel Islands’ protocols.

Parks Canada’s marine policy (Parks This is a historic time of rapid expansion Canada 1994; Mercier and Mondor 1995) both in marine monitoring science (Kramer alludes to goals for a monitoring program 1994; Schmitt and Osenberg 1996) and as follows: governments’ commitments to marine monitoring. In Canada, monitoring marine • ensuring long-term viability of marine environmental quality is an important ecosystems; initiative for both EC and DFO. Marine • understanding natural spatial and monitoring is one of DFO’s three temporal variability in structure (e.g., components of their Ocean Management biodiversity) and function (e.g., Strategy under the Oceans Act. Both EC and production); DFO have collaborated through the • learning how human impacts such as National Marine Indicators Working Group harvest are embedded within the to develop categories of marine indicators - background of natural variability; and parameters tracked over time to reveal trends in processes of interest such as • relating present-day ecosystem resource use or ecosystem health (Smiley et conditions to past ecosystem conditions. al. 1998; Vandermeulen 1998). The Working Group drafted a five-step process for The U.S. National Park Service has been selecting indicators of marine ecosystems as monitoring in marine national parks for >20 follows: years. Davis (1993) summarized the utility

123 • draft as issue statement on reasons for worthy of monitoring. On the other hand, concern within a stress-condition-effect habitat destruction and overharvest are framework; manageable within a conservation area. If • substantiate fully the claims of concern the spatial scales of threats match those of for each issue statement; conservation areas, that should be the prime • explore ecosystem perturbation/ management focus. However, Parks function by linking sections of the Canada marine policy is clear that in order framework, e.g., human activity with to support our regional representation environmental condition; mandate, and in keeping with the openness • screen the long-list of potential and dynamism of marine environments, we indicators into a working short-list; and must be concerned “well beyond” marine • draft summary report on the conservation area boundaries (Parks Canada 1994). For example, we may need deliberations yielding the selected to monitor migratory species (e.g., shore indicators. , whales, salmonids) that occupy Gwaii Haanas only temporarily, but nonetheless DFO has also begun formulating pelagic have important socioeconomic and ecosystem monitoring (Anderson 2001). ecosystem values. Although active migration is not commonly an issue among Parks Canada has not been part of the invertebrates, larvae and pelagic adults can interagency marine monitoring cooperation, be transported over long distances. but this will change in the context of Gwaii Therefore, invertebrate issues outside of Haanas. Fisheries and Oceans Canada has Gwaii Haanas, such as fishing in northern launched a national initiative to facilitate Haida Gwaii, or larval transport from other use of indicators towards marine areas into Gwaii Haanas warrant attention, environmental quality (DFO 2000 a). As perhaps through interagency cooperation. well, the United States has issued its first nation-wide strategy for coastal marine Protected marine areas can also facilitate monitoring (CRMSW 2000 [http:// shellfish stock assessment. In a sea www.cleanwater.gov]). These cucumber diving fishery study in Channel developments underscore the advances in Islands National Park, a fishery- marine monitoring issues that Gwaii independent survey compared stocks in Haanas managers need to be aware of when fished and no-take areas before and after the planning interagency cooperation for onset of fishing (Schroeter et al. 2001). The monitoring decision-making . data revealed that this survey gave more accurate estimates of fished stocks than There also is monitoring in aid of ecosystem data based on fishery-dependent (from the restoration or rehabilitation within a fishery) catch per unit effort data. Thus, conservation area. Monitoring could help Channel Island’s long-term monitoring identify, locate and estimate spatial scale of commitment and enforcement of no-take threats to marine ecosystems. Spatial areas exemplified parks’ potentially decision support tools embedded in GIS are “critically important” roles in stock being developed for monitoring in marine assessment (Schroeter et al. 2001). parks such as the Great Barrier Reef Marine Park (Puotinen 1994). For site management, Without interannual data on distribution we must know which threats can be locally and abundance of species, comparisons managed. Climate change, species between locations may have limited value introduction and external (non-point (Underwood and Kennelly 1990). Using source) pollution cannot, although they are

124 plant communities, although also applicable intertidal and subtidal communities were to communities, these authors among the selected marine environmental proposed the following two stages towards indicators with a suite of measures such as understanding intertidal and subtidal rocky community structure, biodiversity indices, shore communities: recovery of exploited species and behaviour of unexploited species (Rowe et al. 1999). • quantitative studies to describe the Certainly in the last decade, monitoring of patterns of distribution and abundance intertidal invertebrates alone or along with of species with appropriate spatial and seaweeds has become common in national temporal replication to estimate marine parks along the Northeast Pacific variances among and within shores and coast (Table 29). Particularly important for among years and seasons; and use in monitoring has been the persistence • experimental analyses of the processes and dominance on rocky intertidal shores causing the observed patterns. by key epifauna groups such as mussels and barnacles. These groups have well known The latter is the evolutionary end-point of life histories and are amenable to a range of such work and has not yet been achieved in non-destructive monitoring techniques at temperate coastal systems world-wide. fixed sites such as photographing quadrats (squares of known area). However, it can also be argued that monitoring should After clarifying management information include all of the larger animals not just one needs and monitoring objectives, a key or two species, given that each species has a challenge is deciding what marine different ecological niche and may be ecosystem attributes, components or subject to differing stresses. Gwaii Haanas associated species to monitor as discussed already has a baseline on the occurrence of above from the EC/DFO collaboration. conspicuous, dominant intertidal Soliciting the judgement of experts invertebrate epifauna to build upon as (“Delphi” technique) has been used to select Harper et al. (1994) used invertebrate marine invertebrate indicator species from species as well as seaweeds and other ecosystem components by the U.S. National marine plants to discriminate community Park Service (Davis et al. 1994). Further, types from aerial photographs of Gwaii sampling methods must provide unbiased Haanas’ shores (see Table 12). We do not and statistically powerful results while have the same for sediment-dwelling respecting costs and logistics (Gibbs et al. infauna. 1998). Although threat-specific monitoring will be an important part of a comprehensive monitoring program, it is Sediment infauna have been much less used also necessary to develop an integrated for intertidal monitoring. Channel Islands approach to ecosystem monitoring National Park, for example, has a sand (Robinson 2001). beach and lagoon monitoring protocol (Dugan et al. 1990) using six beach invertebrate species ( and Monitoring Invertebrates in Gwaii Haanas mollusk) and three lagoonal (estuarine) non-insect invertebrate species. In the Of all the marine invertebrate habitats in Broken Group Islands unit of Pacific Rim British Columbia, the intertidal has National Park, intertidal burrowing bivalve historically received the most attention populations are monitored to assess levels (Lewis and Quayle 1972; Carefoot 1977). At of beach disturbance (H. Holmes, Pacific a workshop assessing ecological status Rim National Park, personal communication). hosted by Pacific Rim National Park, Although Gwaii Haanas has a coastline

125 126 comprising <10% is sandy shores ( »75% of declining pink abalone (Haliotis corrugata) the shoreline is rocky), sandy/muddy densities revealed that the 15 ha Anacapa habitats should not be overlooked in Island no-take area within Channel Island selecting intertidal species to monitor, National Park was too small to sustain especially as they also represent estuarine historical levels of the species. A potentially habitats. useful monitoring tool for recruitment is deploying samplers for post-larval, settling Subtidal invertebrates are less prominent in invertebrates. These have proved useful in parks monitoring than intertidal monitoring abalone and sea urchin invertebrates. A major exception is Channel settlement (Harrold et al. 1991; Davis 1995). Islands which has been recording 38 A wide range of population parameters of invertebrate species within their kelp forest conspicuous subtidal invertebrates can, monitoring protocol since the early 1980s therefore, be used to monitor efficacy of (Davis et al. 1994). There is, however, a area protection. relatively long history of subtidal long-term trend studies from that are Zacharias and Roff (2001 b) review the ideas applicable to protected area monitoring, but about using particular species in marine have yet to be widely embraced in North conservation management. Parks are ideal America (Lundav 1986; Hiscock 1987). places to establish long-term invertebrate “biomonitoring.” Sessile or sedentary An important source of invertebrate invertebrates, principally bivalves, monitoring data could come from (barnacles, amphipods) and shellfisheries. Shellfisheries will be an polychaetes, have been used as sentinels of integral part of Gwaii Haanas future so that marine environmental quality since the information on target species will be 1960s (Martin and Richardson 1991; important to long-term management. Data Pocklington and Wells 1992). There is an come from both the fisheries themselves extensive literature in the field of and from fisheries-independent surveys biomonitor invertebrates (Rainbow and such as diving stock surveys. It may be that Phillips 1993; Kramer 1994; Schmitt and some commercial species will become Osenberg 1996). These species accumulate sentinel indicators of Gwaii Haanas’ well- pollutants such as metals and hydrocarbons being. In recent research projects, shellfish in their tissues, thus permitting inferences have been useful in demonstrating in-area on the bioavailability of contaminants to effects of protected subtidal areas. For marine ecosystems. Mussels (Mytilus and example, Edgar and Barrett (1997, 1999) Perna spp.) have been particularly widely reported from Tasmania that sea urchin and used (Dame 1996). Gwaii Haanas has numbers and abalone sizes increased abundant mussel populations as well as inside protected areas. Kelly et al. (2000) long-lived geoduck populations that could found that New Zealand lobster densities, function as biomonitors through shell body sizes, biomass and egg production growth parameters (Noakes and Campbell increased inside protected areas. Both 1992). studies were comparisons over time between protected and nearby unprotected In closing, an integrated approach linking areas. In British Columbia, northern offshore and nearshore phenomena will abalone from a protected area increased in characterize the role of invertebrates in density, size and likely reproductive output marine monitoring in Gwaii Haanas. (egg production) over time (Wallace 1999). Robinson (2001) explained that oceanic On the other hand, Davis (2000) found that processes operate at different spatial and

127 time scales influencing the structure and sufficient camping or commercial tour function of Gwaii Haanas’ marine guiding are the main options. Commercial ecosystems. The spatial scales of ocean tour operations account for »70% of visitors climate include local (within kilometers), and »75% of these enjoy multi-day trips. regional (10s to100s of km), coast-wide (100s All visitors attend a mandatory orientation to1,000s of km), and ocean-basin ( »10,000s in which responsible conduct (e.g., no-trace km), while the time scales of ocean climate camping) and respect for the environment include event (weeks), seasonal (months), and Haida culture are stressed. interannual (< 10 years), regime (10s to100s Environmental quality and wilderness of years), and millennial ( »1,000s of years). values such as undisturbed shorelines and For example, the Gwaii Haanas continental solitude are the major expectations of shelf ecosystem report recommended ecotourist visitors. Most of Gwaii Haanas’ monitoring fundamental oceanic variables visitor impacts are coastal due to small boat (Robinson et al. 1999). As well, the Pacific (e.g., kayak) access to the intertidal and Rim workshop (Rowe et al. 1999) yielded back-beach use for camping. Gwaii Haanas’ numerous marine environmental indicators uplands are not commonly visited. such as an “Ocean Climate Index” by assembling remote sensing (satellite) The literature on roles of tourism in marine imagery, meteorological data and assessing conservation has been dominated by studies available physical circulation models. in tropical, coral reef areas (Salm 1985; Robinson (2001) demonstrated that Agardy 1993; Shafer and Benzaken 1998). abundant data from agencies are available Concerns included the paradox of over the World Wide Web concerning large- increasing tourist and recreation impacts scale and relatively long-term changes in after protected status is declared (Jones coastal ocean climate. Most of these data 1994) and, therefore, the need for zoning to are collected at fine time scales (e.g., days) decrease multiple use conflicts in crowded over relatively long time periods. Parks coastal areas (Agardy 1993). The situation Canada has a role of assembling and for Gwaii Haanas is different, as its analyzing these data, and providing remoteness and attendant high costs for interpretations, applicable to all marine visitation limit tourism pressures. But, that groups such as invertebrates, for park is just for now. What directions future management. tourism will take, such as small cruise ships or diving operations, are speculative and Marine Tourism and Visitor Effects on will require planning. The current visitor Intertidal Invertebrates effects on Gwaii Haanas are mainly camping, beach walking, fishing and Gwaii Haanas had »1870 visitors boating. comprising 9,773 visitor-day/nights in 2000. This is among the largest number of The field of “recreation ecology” (Liddle 1991) “backcountry” (wilderness) visitations began with monitoring trampling effects of nation-wide for Parks Canada. There is a visitors to terrestrial parks. Studies have detailed Backcountry Management Plan expanded into trampling effects on (Gajda 1999) for Gwaii Haanas. Visitor intertidal rocky shores (Keough and Quinn databases on GIS have been maintained 1991) and coral reef flats (Hawkins and since 1997 and there is a commitment to Roberts 1993). The full range of visitor long-term campsite monitoring. Because effects on marine conservation areas include Gwaii Haanas is virtually uninhabited and trampling, fishing, diving, boating and off- has little infrastructure for visitors, self-

128 road vehicles as reviewed by McCrone intertidal species for visitor impact monitors (2001). until the natural variation and patchiness of intertidal species are better understood and Certain areas of Gwaii Haanas such as more discernable from variation caused by Dolomite (Burnaby) Narrows at low tide, human impacts. experience appreciable visitor traffic including trampling the intertidal biota Intertidal fishing for subsistence, recreation during visitor season (May to September) (food, bait, curio) and sale have been daytime low tides, as mentioned by Gajda investigated as well. Important rocky (1999). This impact is unquantified, but intertidal studies for small-scale given the adoption of the precautionary (“artisanal”) commercial fishing come from approach in the proposed Canada National Chile (Castilla and Fernandez 1998; Castilla Marine Conservation Areas Act, area closure 1999), indigenous peoples’ gathering from could be implemented based on trampling South Africa (Lasiak 1998), and recreational concerns until data on such impacts are fishing from the U.S. (Addessi 1994) and analyzed. It would be optimal to monitor Australia (Kingsford et al. 1991). Studies on trampling to objectively verify any effects of indigenous peoples’ gathering of ecosystem concerns supporting long-term infauna from intertidal soft sediments area closure. The current management include Australia (Catterall and Poiner recommendation is to promote visitor- 1987) and South Africa (Kyle et al. 1997). vessel float-through rather than intertidal McCrone (2001) summarized findings and walking. concluded that commercial fishing such as in Chile had appreciable direct effects on There is a growing rocky intertidal target species’ abundance and indirect trampling literature in which protected effects on community structure with the seashore areas in Australia (Keough and removal of targeted predators and grazers Quinn 1998), New Zealand (McCrone 2001), (Castilla and Fernendez 1998). Similarly in South Africa (Bally and Griffiths 1989) and South Africa subsistence gatherers lowered the U.S. (Brosnan 1993; Brosnan and abundance of filter-feeding mussels and Crumrine 1994) have been the sites for grazers that increased seaweed cover and assessing visitor effects. Intertidal their attendant invertebrate populations invertebrates are included in these studies (Lasiak 1998). Recreational effects were along with seaweeds; turf-forming species localized, usually within 200 m of beach being the most resistant. Decreases in size access, but nonetheless significantly and density of invertebrates have been reduced some species’ densities and reported. modified aspects of community structure (Addessi 1994). On sediment beaches, Pacific Rim National Park initiated the first molluscs and crustaceans vary in their visitor impact assessments of the rocky vulnerability to fishing impacts according to intertidal in Pacific Canada (Rowe et al. their life-histories and refuges (e.g., 1999). The monitoring protocol, based on burying) from human predation. Catterall the DFO “Shorekeeper’s Manual” (Jamieson et and Poiner (1987) state that for all intertidal al. 1999) is described here in Table 29. Data habitats that life-history and habitat included observations on barnacles, information can enable a priori prediction anemonies and mussels in tide pool, which intertidal populations could be most horizontal bench, crevice and vertical rock susceptible to depletion. In Gwaii Haanas, face substrates. Keough and Quinn (1998), marine recreational fishing is permitted however, stress caution in the use of although there is total closure of all bivalve mollusk species due to the threat of

129 paralytic shellfish poisoning and for interactions affecting adults.” (Roughgarden northern abalone due to low stocks. et al. 1994) Recreational fishing of other groups such as crustaceans (prawn and crab) by trapping is “Many harvested marine invertebrate permitted. populations are metapopulations, composed of relatively sedentary subpopulations connected Motorized vessel traffic in Gwaii Haanas is by dispersing larval stages.” (Botsford et al. from the commercial fishing and tourism 1998) industries and independent visitors. Grey- water (toilet) discharge and fuel spills “Incorporation of selected species dynamics into appear the most likely impacts. MPA rationalization may have to be based on Commercial tour operators have been asked extrapolation from existing data, common sense to limit anchoring at any one location to no and intuition rather than hard scientific longer than three days. The Gwaii Haanas evidence.” (Jamieson and Levings 1998) warden service has recently implemented the MILES + Occurrence Database for “Predicting the consequences of the interaction recording incidents relevant to public safety, between reserve configuration and the law enforcement, environmental protection connectivity pattern is critical to the design of and resource management. In time, this optimally functioning reserves, but prediction will help develop an understanding of the requires detailed information not only on life- actual threats by a wide range of visitor history characteristics and abundance patterns boating activities in Gwaii Haanas. for the target species but also on hydrodynamic current patterns.” (Stockhausen et al. 2000) THE CONTRIBUTION OF SHELLFISHERIES TO MARINE AREA “The greatest potential increases in yield as a MANAGEMENT result of protected areas are achieved with spatially persistent, i.e., relatively sedentary “…. without a knowledge of the ecology of species.” (Jamieson and Levings 2001) larvae an understanding of the benthos is quite impossible.” (Scheltma 1986) It is relevant to analyze how shellfish- associated knowledge may aid consultation “Recent work in marine ecology has reaffirmed towards management of marine area an insight from fisheries science that knowledge conservation. The citations above reveal the about the production, dissemination and success critical importance of larval dynamics to of propagules can guide our management of adult benthic invertebrate populations and populations and assemblages.” (Fairweather underscore perhaps the core science-based 1991) issue confronting marine area conservation. The art of marine area conservation for “Appropriately designed fishery refugia may be Parks Canada and its partners in Gwaii one of the tools used to address the management Haanas will be to manage multiple and rehabilitation of coastal stocks and sustainable uses, such as commercial ecosystems in the next century.” (Dugan and fishing, while maintaining long-term Davis 1993a) ecosystem structure and function. Indeed, if the proposed Canada National Marine “In coastal marine systems, physical Conservation Areas Act is passed, oceanographic processes affecting larval stages management “…without compromising the are as, or more important than, biological structure and function of the ecosystems” [section 4(4)] would be the law – not an

130 option. Further, this varies from the Oceans adaptive shellfish management is being Act that, although supporting the ecosystem applied to sea cucumber management on a approach in its preamble, does not explicitly trial basis as well as to new and developing mention ecosystem structure and function fisheries (B. Adkins, DFO, personal in its section 35 on marine protected area communication). management criteria. Secondly, therefore, the challenge will also be to observe and Zoning, Fishing and Refugia balance the legislated mandates of other agencies (e.g., DFO, EC, NRCan) within “……management for direct use (e.g., fisheries), marine conservation areas. for indirect use (e.g., heritage and existence values), and for ensuring protection of essential Overall, fisheries science can contribute ecosystem services, ultimately must be much in support of the broader context of accomplished through zoning, which requires marine conservation (Fairweather 1991; designating different areas to meet different Jamieson and Levings 2001). Conversely, goals.” (NRC 2001) promoting the potential contribution of protected marine areas to fisheries research “Renewable resource conservation through has been a recurrent theme from the protected areas may imply some probable beginning of the technical (non-fisheries) restriction of human fishing activity but is likely marine conservation literature (Wallis 1958), to enhance achievable yields in adjacent and remains an important consideration exploited areas.” (Jamieson and Levings (Levings and Jamieson 1999). Further, 2001) much has been written about marine conservation areas as instruments of Coexisting ecologically sustainable uses are fisheries management including as enabled by zoning. Zoning includes the “insurance” against effects of over-fishing notion of refugia (“no-take” areas), which (Allison et al. 1998; Guenette et al. 1998; are a vibrant topic in coastal fisheries and Lauck et al. 1998; NRC 1999, 2001). The key marine conservation thinking (Dugan and point is that, through appropriate Davis 1993a,b; Botsford et al. 1997; Roberts knowledge-based management, protected 1998, 2000; Hastings and Botsford 1999; marine areas can be an asset to nearby Murray et al. 1999; NRC 1999, 2001; Dayton fisheries. et al. 2000; Sumaila et al. 2000; Roberts et al. 2001; Sloan 2002). Gwaii Haanas could be useful for exploring adaptive shellfisheries management. Shellfish issues will contribute to defining Results could provide feed-back enabling Parks Canada’s policy commitment to managers to learn-by-doing and alter zoning in marine conservation areas such as management to achieve desired shellfishery Gwaii Haanas (Parks Canada 1994). The performance criteria. Orensanz and shellfish species in Haida Gwaii with Jamieson (1998) suggested that creation of benthic adults comprise local sub- marine conservation area networks should populations connected by planktonic larval employ adaptive shellfish management to dispersal. That is, benthic adult shellfish learn from the natural system’s responses. populations are relatively “spatially Adaptive management is still not being persistent” (Orensanz and Jamieson 1998; applied widely in marine conservation Jamieson and Levings 2001). Recruitment is (Walters 1997; Parma et al 1998), although it meant here, not in the strict fishery sense, is gaining acceptance (Jamieson and but in the broad sense of addition of new Levings 2001). In British Columbia, individuals to populations (Caley et al.

131 1996). Of course, besides sound science, is the relative importance of recruitment achieving zoning must be done through the versus post-recruitment processes in commitment to appropriate public determining population size and consultation – accepted as critical to success structure, i.e., to what extent does in marine conservation (Sumaila et al. 2000). population abundance depend on recruitment (establishment of initial On the science side, sizes of refugia depend, population pattern) compared to in part, on the dispersal ability of species or competition/predation/facilitation/ species groups targeted for conservation. disturbance (modifiers of population Their size may also depend on how well pattern). they are buffered from outside impacts. Which areas are “sources” (contribute The implication is that, for years to come, disproportionately large numbers of our scientific understanding of processes recruits) and which are “sinks” (receive fundamental to marine conservation will recruits but contribute little) of recruits is lag behind the pressing need for critical management information (Ogden implementation. Hence, the important role 1997; Ballantine 1997 a,b; Jamieson and of the precautionary approach. Ludwig et al. Levings 1998; Roberts 1998, 2000; Roberts et (1993) were among the first to champion not al. 2001). The Great Barrier Reef Marine waiting for science consensus before Park, for example, has for some time looked creating marine reserves as a common-sense at how larval dispersal modeling can precautionary measure. We should be support zoning (James et al. 1990). Botsford guided by the precautionary approach and et al. (1994) warn, for example, that spatially an ecosystem viewpoint (both Parks Canada managed populations of red sea urchins and DFO commitments) blended with an would go extinct if distances between adaptive management ethic, what science refugia exceed the scale of larval dispersal. we do know and “common sense and intuition” (Jamieson and Levings 1998). It is ironic that the core marine area conservation connectivity (“source/sink”) On the socioeconomic side, the unique issue should also be widely recognized as rights of First Nations and the interests of perhaps the greatest unsolved question in other stakeholders such as the commercial marine biology – that is, the degree of fishery sector, coastal communities, connectedness between local populations government agencies and NGOs must blend (Scheltema 1986; Gaines and Bertness 1992; to reconcile the complexities of Caley et al. 1996; Hunt and Scheibling 1997). conservation, commerce and culture. Caley et al. (1996) summarized the basic unresolved science issues in recruitment as Insights Provided by Shellfish Species follows: “Given the sedentary life-history characteristics • poor understanding of why recruitment of invertebrates and the nature of their fishing varies in space and time; process, management measures that explicitly • limited knowledge on the extent to acknowledge spatial structure are most which spatial and temporal variation in suitable.” (Orensanz and Jamieson 1998) recruitment influence variation in adult populations; and Biology combined with commerce • perhaps the most difficult and intensifies focus on the roles of commercial important gap in understanding benthic species in marine area conservation. Yet, and demersal (near-bottom) populations from the 1995 North Pacific Symposium on

132 Invertebrate Stock Assessment and cosmopolitan (Pacific and Atlantic) Management in Nanaimo, Orensanz and distribution. Larval dispersal distances Jamieson (1998) concluded that range from settlement near the parental “Conservation issues may have been the most stock (Young and Chia 1982) to settlement significantly underrepresented topic at this more than 2,000 km away from spawning Symposium.” In other words, these are early adults (Booth and Phillips 1994). Durations days for implementing linkages between of planktonic larval phases range from two shellfisheries management and marine area hours (Strathmann 1987 - tunicates) to two conservation beyond single species area years (Booth 1994 – spiny ). Finally, closures. The spatial persistence of many some invertebrates, such as many snail benthic marine invertebrates makes them species, have no free-living larval stage. good study subjects for conserving marine areas. A sound approach to science-based As examples of this variability among management includes characterization of Haida Gwaii shellfish species, Table 30 lists potential zones with their associated biological characteristics of five species with invertebrate species. This could put Gwaii benthic adults. Just among these species, Haanas at the forefront of marine benthic adult life-history and planktonic conservation science in Canada. larval life-history vary dramatically. These are just five among the thousands of marine Patchy adult shellfish populations invertebrate species in the Haida Gwaii associated with identifiable habitat or region! ecosystems types, such as geoduck in sand beds or northern abalone on kelp-forested At one end of the spectrum is northern rock reefs, facilitate potential zone abalone with the following characteristics: demarcation. Further, the dispersal capabilities of these species’ larvae provide • sedentary adults that form pre- insights into scale and connectedness for spawning clusters for synchronous planning zone numbers, sizes, shapes and spawning; locations. Having said that, be warned that • external fertilization of broadcasted both the invertebrate species’ biology and gametes for which fertilization nearshore oceanography in Gwaii Haanas efficiency declines with a decline in are poorly known. Therefore, blending adult densities; and biology and oceanography in conservation • short-duration, non-feeding larvae area development will long be a work-in- unlikely to travel far before settling. progress. Therefore, the patch size, location and There is immense variety in the early life- spacing of adult populations is critical. history traits among marine invertebrates with benthic adults (Scheltema 1986). At the other end of the spectrum is Looking at the larval lives of invertebrates Dungeness crab with the following is important. Invertebrates tend to have characteristics: more limited dispersal potential than fishes and planktonic larval duration is often, but not necessarily, related to the geographic • semi-sedentary adult males (tagged crab range of adult invertebrates (Bradbury and can move >60 km [I. Winther, DFO, Snelgrove 2001). A good example is the personal communication]) which pair and brittle star Amphipholis squamata that bears copulate with post-moult (soft-shelled) live young (i.e., no larval phase), yet has a females;

133 • sperm retention for subsequent fertilization and fertilized egg retention by brooding females until larvae hatch; and • long-lived, feeding larvae that travel 100s to 1,000s of km and mix broadly in offshore waters before returning inshore to settle, perhaps very distant from their location of hatching.

There is not a clear linkage of the densities of adult Dungeness crab populations with the production of larvae by those populations. It is possible, however, to model the influence of near-shore current patterns on spatial distribution of Dungeness crab recruitment at a scale of 100s of km (Crawford and Jamieson 1996; Wing et al. 1995, 1998).

Figure 33 illustrates likely applicable models of northern abalone and Dungeness crab larval replenishment adapted from Carr and Reed (1993). These models assume reef-dwelling adult populations, which Dungeness crab is not, but the illustration nonetheless summarizes the Dungeness crab situation. The two

134