[From the U.S. Government Printing Office, www.gpo.gov]



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                              1994
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                                                                 114
                                                                                                                              XI




                                                                                  Alaska Sea Grant
                                                                                  College Program
                                                                                          laska Fairbanks
                                                                           University of A
                                                                                   P.O. Box 755040
                                                                                                                             14"A
                                                                            Fairbanks, Alaska 99775-5040
                                  f"N'
                                                                                   (907) 474-6707
                                                                                  Fax (907) 474-6285
                                                 @'Iy 10
                                                                      A,                                    IV.
                                                                                  AK-SG-94-01   $3.00









                                           Elmer E. Rasmuson Library Cataloging-in-Publication Data

                                           Wyllie-Echeverria, S.
                                                Managing seagrass systems in western North America: research gaps and
                                           needs / S. Wyllie-Echeverria, R.M. Thom.

                                                "AK-SC-94-Ol."
                                                Includes bibliographical references.
                                                1. Seagrasses-North America-Pacific Coast. I. Thom, Ronald M. 11. Alaska
                                           Sea Grant College Program. III. Title. IV. Series: Alaska Sea Grant report; no.
                                           94-01.


                                           QK495A14W95 1994




                                           ISBN 1-56612-022-5








                                           CREDITS

                                           Cover design is by Susan Burroughs, and background photo on cover is by Sandy
                                           Wyllie-Echeverria (photo has been altered). Editing is by Sue Keller and Carol
                                           Kaynor. Layout is by Carol Kaynor.
                                                This book is the result of work sponsored by the University of Alaska Sea Grant
                                           College Program, which is cooperatively supported by the U.S. Department of
                                           Commerce, NOAA Office of Sea Grant and Extramural Programs, under grant no.
                                           NA90AA-D-SG066, projects M/81-01 and A/75-01; and by the University of Alaska
                                           with state funds. The University of Alaska is an affirmative action/equal opportunity
                                           employer and educational institution.
                                                Sea Grant is a unique partnership with public and private sectors combining
                                           research, education, and technology transfer for public service. This national
                                           network of universities meets changing environmental and economic needs of
                                           people in our coastal, ocean, and Great Lakes regions.


                                           Property Of                           J1, I.
                               40AA Coastal Services Center
                                               Library






                                              Contents



                                                 The Authors                                                           iv
                                                 Acknowledgments                                                       iv
                                                 Dedication                                                            iv


                                              Introduction                                                             I

                                              History of the West Coast Seagrass Group                                 3
                                                 California Efforts                                                    3
                                                 Washington Efforts                                                    5
                                                 Pacific Coast Seagrass Newsletter                                     7
                                                 Monterey Symposium                                                    7

                                              Seagrass Research Needs on the West Coast                                7
                                                 Physiology                                                            8
                                                 Population Dynamics and Reproduction                                  9
                                                 Environmental Studies                                                 12

                                              Concluding Remarks                                                       14

                                              References                                                               is

                                              Appendix                                                                 16
                                                 Seagrass Research in Southwestern British
                                                 Columbia, Canada                                                      16
                                                 Seagrass Research in Baja California, Mexico                          18

                                              Contributors                                                             21









                                                                                   id








                                             The Authors

                                             Sandy Wyllie-Echeverria is a Visiting Scholar at the School of Marine Affairs,
                                             University of Washington, where he is a member of the Interdisciplinary Seagrass
                                             Working Group (ISWG). The ISWG formed in fall 1992 to examine seagrass manage-
                                             ment programs in the Pacific Northwest using an interdisciplinary approach. Wyllie-
                                             Echeverria's research interests include seagrass population dynamics, restoration of
                                             damaged seagrass systems, subtidal seagrass mapping techniques, effects of light
                                             environments on seagrass distribution,. history of seagrass science and policy, and
                                             traditional use of seagrass resources. For most of the last decade, at several locations
                                             along the West Coast of North America, he has given public lectures, participated in
                                             university classes, and made presentations at scientific meetings in an effort to craft a
                                             policy toward sound management of the seagrasses of the Northeast Pacific. He
                                             initiated the work for this booklet while a graduate student at the University of Alaska
                                             Fairbanks.
                                                  Dr. Ronald M. Thom is a Senior Research Scientist at Battelle's Marine Sciences
                                             Laboratory in Sequim, Washington. Prior to moving to Battelle in 1990, Thom was a
                                             researcher in the Fisheries Research Institute at the University of Washington. His 21
                                             years of professional experience include employment as a biologist for Los Angeles
                                             County and the Seattle District Corps of Engineers. His research interests include
                                             benthic primary production, the effects of pollution on nearshore marine communi-
                                             ties, habitat restoration, the effects of climate change on nearshore systems, the
                                             ecology of fisheries resources in nearshore systems, and biodegradation of petroleum.
                                             Most of his work has been conducted in Puget Sound, southern California, and
                                             Alaska. Thom has served on numerous committees, including a five-year appoint-
                                             ment as Chair of the Technical Advisory Committee to the Puget Sound Estuary
                                             Program.

                                             Acknowledgments
                                             The authors gratefully acknowledge the Western Society of Naturalists for permitting
                                             us to hold the West Coast Seagrass Symposium in conjunction with WSN's 1990
                                             annual meeting. We thank Alaska Sea Grant Director Ron Dearborn for his ideas,
                                             which helped frame the symposium that formed the basis for this report, and for
                                             providing staff support-the editorial work by Sue Keller and Carol Kaynor was key
                                             in the development of the final product. We also appreciate the thoughtful and
                                             thorough reviews of R.S. Alberte, J.T. Carlton, and PG. Harrison. Blake Feist provided
                                             the map depicting seagra5s distribution. Finally, we acknowledge the help of our
                                             respective families, whose patience and support during this project was invaluable.


                                             Dedication

                                             This booklet is dedicated to R.C. Phillips, whose pioneering seagrass research has
                                             been an inspiration to all seagrass scientists. Dr. Phillips continues to be active in
                                             seagrass research from his post at Battelle-Pacific Northwest Laboratory in Richland,
                                             Washington.



                                                                                        iv










                                      Introduction

                                      This report summarizes the history and direction of efforts to connect
                                      scientists and resource managers along the West Coast of North America,
                                      to develop a regional seagrass management plan. These activities began in
                                      the 1980s and reached a plateau at the West Coast Seagrass Symposium in
                                      December 1990. Second, it summarizes seagrass research gaps and needs
                                      in this region. Information is derived from research priorities identified by
                                      two seagrass working groups and by seagrass scientists in this region (see
                                      Contributors).
                                           Six species of temperate and cold-water seagrasses grow on the West
                                      Coast (Figure 1). All species belong to the family Potornogetonaceae but
                                      are differentiated into two genera: Zostera and Phyllospadix (Phillips and
                                      Menez 1988). The genus Zostera has three species: Z. asiatica, Z. japonica,
                                      and Z. matina, eelgrass. The genus Phyllospadix, surfgrass, also has three
                                      species: P serrulatus, R scouteri, and P torreyi. Most probably, four of the
                                      species (Z. mafina, R serrulatus, P scouleri, and R torreyi) have been
                                      growing along the rocky shores and soft-bottom habitats of this region
                                      since the Pliocene (Phillips and Menez 1988, Domning 1977). Z. japonica
                                      probably was introduced in the 1930s and early 1940s (Harrison and Bigley
                                      1982), while Z. asiatica was only recently described in this region (Phillips
                                      and Wyllie-Echeverria 1990). In some accounts the brackish water plant
                                      Ruppia maritima is classified as a seagrass (e.g., den Hartog 1970). This
                                      plant grows in western North America (Mason 1957, Steward et al. 1963,
                                      Brayshaw 1985); however, it has not often been considered a seagrass in
                                      this region by either research science or resource management and is not
                                      referenced in this report.
                                           Development pressure in the coastal habitats of the world's oceans is
                                      intense, and seagrasses are often impacted by resulting coastal modifica-
                                      tion. Along the West Coast, it is common for developers to be dismayed at
                                      the disarray of opinion on the relative ecological importance of seagrass
                                      meadows, factors that may cause significant injury to the meadows, and
                                      the possibility of restoring impacted meadows or creating new ones. This






                                        2 Managing Seagrass Systems in Western North America








                                                                        Alaska
                                                                                                       V
                                                                                                                _4"





                                                                                                CANADA






                                                                                                  C,




                                                                                                           46
                                                                                                       U.S.








                                                                                                                 4-1
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                                        Figure 1. Seagrass distribution in the temperate waters of the Northeast Pacific
                                                 (Phillips and Menez 1988, Phillips and Wyllie-Echeverria 1990). Symbols
                                                 represent general ranges.






                                                                                      Research Gaps and Needs 3


                                     disarray is a function of several phenomena. Most notable are (1) lack of
                                     data on some fundamental questions concerning seagrass biology, autecol-
                                     ogy, and community structure; (2) ineffective communication of the
                                     reliable scientific information that does exist; and (3) the difficulty of
                                     making reasonable, rational decisions in resource management and
                                     development. Investigators in turn are concerned because the manage-
                                     ment of seagrass systems is carried out in the absence of adequate scien-
                                     tific information. In the final analysis, developers, resource managers, and
                                     scientists feel powerless to chart a course toward sound management.
                                         Yet a spirit of cooperation exists among researchers studying seagrass
                                     on the West Coast. This cooperative spirit should lend itself to the develop-
                                     ment of coordinated research projects along the entire coast, encompass-
                                     ing long time-series data sets and collaborative investigations. In addition,
                                     there is intense interest in developing a better understanding of the
                                     importance of seagrass to fisheries resources.

                                     History of the West Coast Seagrass Group

                                     lWo seagrass working groups formed in the 1980s in California and
                                     Washington, and two regional meetings organized by working group
                                     members were held. These activities are described below.


                                     CALIFORNIA EFFORTS

                                     William Albert Setchell began seagrass studies while at the University of
                                     California, Berkeley, in the 1920s. Setchell (1929) was able to establish an
                                     informal network to facilitate his studies on eelgrass. Although he had two
                                     study sites on the shores of San Francisco Bay, he relied on contributions
                                     from individuals in several states to complete his analyses. Setchell's work
                                     was thorough and comprehensive, and his studies provided a baseline for
                                     other seagrass work in the United States. Unfortunately, this promising
                                     beginning did not provide strong impetus for continued seagrass research
                                     in San Francisco Bay, where research activities have resumed only recently
                                     (Nichols and Pamatmat 1988, Zimmerman et al. 1991, Kitting and Wyllie-
                                     Echeverria 1992).
                                         In 1984, an eelgrass transplant was initiated in San Francisco Bay. The
                                     transplant team reported that lack of data on local eelgrass autecology,






                                     4 Managing Seagrass Systems in Western North America


                                     coupled with water quality conditions, limited transplant success (Fredette
                                     et al. 1988). The team recommended that eelgrass studies be initiated and
                                     maintained and that issues of water quality be addressed. In response to
                                     these recommendations, an interdisciplinary group of scientists and
                                     managers was formed in 1985 to outline eelgrass research priorities for
                                     San Francisco Bay. This group, known as the "ad hoc eelgrass working
                                     group," identified the following priorities:
                                       ï¿½  New and updated surveys to determine the distribution and density of
                                          eelgrass.
                                       ï¿½  Field research to determine the value of eelgrass as a habitat in San
                                          Francisco Bay.
                                       ï¿½  Field research and laboratory studies to determine the physical
                                          parameters that might limit plant distribution.
                                       ï¿½  Detailed investigations of the biology and autecology of eelgrass in San
                                          Francisco Bay.
                                       ï¿½  Continued studies to evaluate effective transplant techniques.
                                          The working group met for two years and continued to discuss
                                     possible research projects and mechanisms for funding. In May 1987, the
                                     first California Eelgrass Symposium was held at the Paul F. Romberg
                                     Tiburon Center for Environmental Studies (RTCES), San Francisco State
                                     University, Tiburon, California. An initial report of resource inventory
                                     results was presented and it was agreed that a survey of San Francisco Bay
                                     eelgrass should become the top research priority. The National Marine
                                     Fisheries Service, Southwest Region, agreed to provide limited funding,
                                     and a survey project began in the summer of 1987. The project had two
                                     objectives: (1) provide a map depicting the geographic range and distribu-
                                     tion of eelgrass in San Francisco Bay, and (2) provide estimates of total
                                     acreage at individual sites (Wyllie-Echeverria 1990, Wyllie-Echeverria and
                                     Rutten 1989).
                                          Commensurate with the third research priority, a research team from
                                     the University of Chicago stationed at Hopkins Marine Station, Pacific
                                     Grove, California, and RTCES, was funded to assess the environmental
                                     suitability of San Francisco Bay for eelgrass. The objectives of the study
                                     were to: (1) determine the relationship between turbidity and light
                                     available for eelgrass growth and reproduction, and (2) determine the






                                                                                     Research Gaps and Needs 5


                                     effect of this light environment on the depth distribution of eelgrass in the
                                     bay (Zimmerman et al. 1991).
                                         During the first eelgrass symposium in 1987, a panel of resource
                                     managers concluded that the symposium format should be continued but
                                     that wider geographic participation should be encouraged. Accordingly, a
                                     second California Eelgrass Symposium, organized by Pacific Southwest
                                     Biological Services, was held in Chula Vista, California, in May 1988.
                                     Symposium participants came from British Columbia, Washington,
                                     California, and Mexico. Speakers addressed research and management
                                     priorities identified at the Tiburon symposium. The issues, summarized in
                                     Merkel and Hoffman (1990), include: (1) case studies from experimental
                                     and compensatory transplants in this region, (2) descriptions of the
                                     contribution of eelgrass to coastal food web dynamics, and (3) discussion of
                                     the primary factors limiting eelgrass distribution and abundance.

                                     WASHINGTON EFFORTS
                                     In autumn 1986, the Washington Department of Natural Resources
                                     (WDNR) invited researchers and resource managers to join in a coopera-
                                     tive effort to evaluate the functional value of eelgrass. At the first meeting,
                                     Thomas Mumford, group coordinator from WDNR, explained that federal,
                                     state, and local decision-makers needed scientifically valid information on
                                     site-specific and regional functional values of seagrass systems. Urgently
                                     needed were (1) maps of the distribution of seagrasses, (2) identification of
                                     the functional values of seagrass systems, and (3) methods for mitigating
                                     loss of functional values. It was recognized that there was growing pres-
                                     sure on seagrass systems and a lack of understanding of the importance of
                                     these systems to natural resource management.
                                         The group met several times in 1987 and developed a list of functional
                                     attributes characteristic of seagrass systems. The topics on the list, which
                                     included physical, biochemical, and biological functions as well as manage-
                                     ment factors, were developed into a five-phase research approach designed
                                     to systematically acquire information needed by decision-makers. The
                                     priorities were:
                                      * Aerial photographic analysis to map seagrass distributions.
                                      0 Subsample surveys to select representative patches for further
                                         research.






                                       6 Managing Seagrass Systems in Western North America


                                        ï¿½  In-depth studies on plant biology, faunal and floral structure, carbon
                                           energetics, and mechanics.
                                        ï¿½  Studies to optimize transplant techniques and assess the functionality
                                           of constructed seagrass systems.
                                        ï¿½  Data synthesis to provide information to decision-makers in a form
                                           that would maximize their ability to manage the resources.
                                           The work of the group was not published, and documentation remains
                                       in the form of meeting notes. Although funding for conducting the
                                       outlined research was not made available, the first phase of the study plan
                                       was partially accomplished by another effort, the Puget Sound Monitoring
                                       Plan (PSMP). PSMP, part of the Puget Sound Estuary Program and partially
                                       funded by the Environmental Protection Agency, resulted in the develop-
                                       ment and implementation of a nearshore habitat monitoring program that
                                       included seagrass meadows (Puget Sound Water Quality Authority 1989).
                                       The program has collected distribution data on eelgrass in Puget Sound
                                       since 1988 and now has the beginnings of a comprehensive database on
                                       seagrass distribution in that area. Data and summaries of the studies are
                                       available from Thomas Mumford, Aquatic Lands Division, Department of
                                       Natural Resources, P.O. Box 42027, Olympia, WA 98504-7027. Data from
                                       Washington other than Puget Sound are largely lacking.
                                           Management of eelgrass in Washington continues to be a major
                                       concern, and the Washington State Department of Fisheries (WDF) formed
                                       in 1989 an intra-agency working group to define eelgrass policy. This group
                                       has met periodically but to date has not published a formal statement
                                       regarding eelgrass resource research or management needs. At present, the
                                       WDF seagrass working group is the most active entity pursuing eelgrass
                                       issues in the state with regard to resources and management.
                                           Development impacts on eelgrass and mitigation of these impacts
                                       currently are the most pressing environmental issues facing the U.S. Army
                                       Corps of Engineers, Seattle District, as well as the regional shipping ports.
                                       Approximately $100 million in development projects have been stopped or
                                       stalled in Washington State from 1990 to 1993 because of these issues (R.
                                       Thom, unpublished data). A perception that eelgrass cannot be successfully
                                       transplanted as mitigation has largely been responsible for denial of
                                       development permits (Thom 1990).






                                                                                           Research Gaps and Needs 7


                                        PACIFIC COAST Swims NEWSLETTER
                                        Production of the Paciflc Coast Seagrass Newsletter was an effort to merge
                                        the goals of the California and Washington seagrass groups and develop a
                                        more regional approach. After several months of discussion and a commit-
                                        ment by the University of Washington Fisheries Research Institute (FRI),
                                        the first issue was mailed in spring 1990.
                                            The newsletter had three goals: to announce meetings and workshops
                                        on seagrass-related topics, to communicate information between seagrass
                                        scientists and resource managers on a regional scale, and to provide a link
                                        between seagrass scientists and resource managers along the West Coast,
                                        including Mexico and Canada. Reasons for focusing on the West Coast
                                        were: (1) links between seagrass scientists and resource managers, al-
                                        though established locally, were not well developed on a regional scale; (2)
                                        much of the funding for seagrass research seemed concentrated in the
                                        eastern United States, and the newsletter editors believed that more
                                        attention should be focused on West Coast systems; and (3) limited
                                        resources demanded focusing efforts on one region.
                                             In 1992, the Pacific Estuarine Research Society (PERS), a regional
                                        society of the Estuarine Research Federation, requested that we consider
                                        reformatting the newsletter as a column in the PERS newsletter. The
                                        column continues to fulfill the goals of the original seagrass newsletter.

                                        MONTEREY SYMPOSIUM
                                        With a small budget from Alaska Sea Grant, a seagrass symposium was
                                        held as part of the annual meeting of the Western Society of Naturalists
                                        (WSN) in December 1990 in Monterey, California. The goal was to clarify
                                        previously identified research questions and suggest regional research
                                        priorities; therefore, invited participants focused on issues of general
                                        concern to the region. Approximately 25 individuals attended and 12
                                        papers were presented.

                                        Seagrass Research Needs on the West Coast
                                        In an effort to capture the insights of West Coast seagrass scientists, the
                                        authors of this report asked each participant in the Monterey seagrass
                                        symposium to provide a list of knowledge gaps and research priorities. We






                                       8 Managing Seagrass Systems in Western North America


                                       also requested contributions from researchers who were unable to attend
                                       the symposium. We have synthesized the remarks of this body of scientists
                                       (see Contributors), whose primary research focus has been the seagrasses
                                       of the West Coast of North America. The intent of this summary is to
                                       inspire new research proposals and activities which, in turn, should guide
                                       seagrass policies and management in the next century.

                                       PHYSIOLOGY

                                       A thorough understanding of seagrass physiology is essential to fully
                                       comprehend the mechanisms by which seagrasses persist in existing
                                       meadows. This knowledge may also facilitate the development of effective
                                       transplant techniques throughout the region. Important research topics
                                       include: thermal requirements and tolerances, light requirements, nutri-
                                       ent requirements, grazer interactions, carbon and nitrogen cycling,
                                       sediment requirements, salinity regimes, and environmental toxicology.
                                       Knowledge of how these factors affect or interact with the maintenance of
                                       viable, healthy, and functional seagrass systems is required. Although
                                       physiological studies have been done for eelgrass, these need to be ex-
                                       panded. In addition, this work should be initiated on the other five species
                                       in the region. There is a vast array of literature on seagrass physiology in
                                       other regions of the world, and this literature and research can provide a
                                       foundation for West Coast studies. However, the distribution of
                                       Phyllospadix torreyi, R scouleti, and R serrulatus is limited to the West
                                       Coast of North America, and Zosterajaponica and Z. asiatica grow only in
                                       temperate Pacific waters. Basic research to describe the physiological
                                       expressions of these plants is necessary before complete and comprehen-
                                       sive understanding will be achieved. Research on the following topics is
                                       critically needed:

                                         Root and shoot nutrient physiology
                                       It is established that seagrasses can assimilate inorganic nutrients through
                                       both roots and leaves. However, the factors controlling uptake rates and
                                       processes are not well established. Interactions between nutrient availabil-
                                       ity in the water column and sediment and uptake rates by leaves and roots
                                       are not understood for all species. Information on this topic is critical to
                                       understand the effects of eutrophication and nutrient limitation. In






                                                                                           Research Gaps and Needs 9


                                        addition, this information would help establish guidelines for optimal
                                        transplanting.

                                        * Carbon and nitrogen allocation and nutrient pool size
                                        Conversion of inorganic nutrients and carbon to organic compounds, and
                                        the storage, transport, and utilization of these compounds, is not well
                                        known. In particular, carbon and nitrogen sinks within vegetative and
                                        reproductive shoots as well as species-specific carbon and nitrogen
                                        distributions have not been established for all species. An understanding of
                                        nutrient pool sizes and the physical, chemical, and biological controls on
                                        interstitial nutrient pool size and composition is needed to describe
                                        pathways of energy allocation and storage. These data also have implica-
                                        tions for modes of energy transfer to coastal food webs.

                                          Interactions of nutrients and light
                                        Human activities resulting in a reduction of light energy in aquatic
                                        systems (e.g., eutrophication and suspended sediment from dredging and
                                        filling) can result in large-scale declines of seagrasses. Studies are needed
                                        to sort out the relationships between epiphytes stimulated by nutrient
                                        additions and the seagrasses to which they are attached. Overabundant
                                        epiphyte loads can cause severe reductions in seagrass growth. This
                                        phenomenon, coupled with declines from reduced light environments,
                                        gives cause for concern.

                                        POPULATION DYNAMICS AND REPRODUCTION

                                        The population dynamics of all species occurring on the West Coast are
                                        poorly understood. Research is needed to describe the similarities and
                                        differences between small, fragmented patches and continuous stands.
                                        Studies should also examine genetic structure and diversity and relate
                                        these features to performance and ecological success. Although recent
                                        studies have clarified seagrass breeding systems, more work is necessary.
                                        Work is especially needed to determine possible effects of human-induced
                                        disturbance (e.g., non-point source contamination resulting from agricul-
                                        ture, industry, logging, and urbanization) on pollination systems and
                                        mechanisms. Finally, detailed maps describing distributions and character-
                                        izing populations are needed for all species throughout the region.






                                     10 Managing Seagrass Systems in Western North Arnerica


                                     Although some maps exist for eelgrass, information for the other species is
                                     very limited.

                                     e Density-dependent features
                                     Currently no research allows us to determine the relationship between
                                     shoot density, patch size, and modifications in substrata and sediment
                                     biogeochemistry. This information is necessary to judge the point at which
                                     a small patch begins to function as a seagrass system, and it has direct
                                     consequences on proper management. For example, guidelines in Wash-
                                     ington now state that a patch of eelgrass with a density of 50 shoots per ml
                                     is a functioning patch. This criterion is based on qualitative observations of
                                     the persistence of patches over several years. Three problems arise from
                                     this type of policy: (1) Before the policy can be adopted in other locations
                                     along the West Coast, adequate field tests at several locations are neces-
                                     sary; (2) criteria should be based on quantitative as well as qualitative
                                     analysis; and (3) although persistence is an ecologically sound measure-
                                     ment of functionality, other parameters such as modified sediment
                                     processes, higher infauna and epifauna densities, and enhanced organic
                                     composition of the sediments are also important indices.


                                       Turnover rates

                                     To date, we have no data to determine the fundamental population param-
                                     eters of birth rate, death rate, and longevity for seagrass populations on the
                                     West Coast. There is also a need to determine the nutrient requirements of
                                     seedlings and to clarify their role in structuring patch size. In addition,
                                     there are no data to document the life expectancy of rhizome tissues or
                                     short shoot tissues. Finally, there is insufficient evidence to conclude
                                     whether plants in small patches are replaced more rapidly than plants in
                                     large, continuous stands. Without a basic understanding of these popula-
                                     tion parameters, there is no rigorous way to evaluate limiting factors or to
                                     estimate the true carrying capacity of a system for a seagrass species.

                                     * Role of genetic diversity and gene flow within populations
                                     It is generally known that higher diversity leads to stability in natural
                                     systems. This is undoubtedly true for seagrass systems; nevertheless, we
                                     have only recently begun to understand the role of genetic diversity in






                                                                                     Research Gaps and Needs 11


                                      maintaining seagrass populations. Major questions to be answered include:
                                      (1) Is disease resistance dependent on increased diversity? (2) Are certain
                                      genetic strains more tolerant of particular disturbance events (e.g., low
                                      light resulting from increases in suspended sediment)? (3) What are the
                                      rates and processes of gene flow between disjunct populations? (4) Does
                                      increased genetic diversity render a population more resilient to invasion
                                      by exotic species? and (5) Do seagrass meadows with higher genetic
                                      diversity offer enhanced ecological support to animal populations?

                                       Pollination studies
                                      Real progress has been made in the last decade in the understanding of
                                      seagrass pollination biology, and this research has collapsed the notion
                                      that seagrasses are largely asexual or rely on crude or inefficient pollina-
                                      tion mechanisms. Research questions that warrant further attention
                                      include: (1) exploration of the physiological mechanism involved in pollen-
                                      stigma interactions, (2) clarification of the relationship between tidal and
                                      current activity and the timing of pollen and seed dispersal, and (3)
                                      elucidation of the relationship between seagrass morphology, pollination
                                      systems, and population structure.

                                        Distribution maps
                                      Several human activities threaten the growth and survival of seagrass in
                                      this region, including dredging and filling, industrial and agricultural
                                      runoff, and erosion from increased logging. Before we can begin to assess
                                      impacts, however, we need a working knowledge of the amount of seagrass
                                      present in the region as a whole. On this score we have little data. Al-
                                      though seagrass resources have been mapped at a few locations, these
                                      maps are usually one-time efforts and involve only one species, eelgrass.
                                      Some efforts have been made to map Z. japonica, but these activities are
                                      very localized. Phyllospadix spp. and Z. asiatica have received less atten
                                      tion. In sum, we have limited knowledge of the regional distribution and
                                      -abundance of all species and are thus unable to assess the relative and
                                      cumulative impact of human activities over time.
                                          Given the continued pressure to develop the margins between land
                                      and sea, accurate and comprehensive knowledge of seagrass distribution
                                      and abundance is essential for prudent and wise management of the






                                    12 Managing Seagrass Systems in Western North America


                                    resource. It is no longer necessary to validate the relative worth of sea-
                                    grasses within the context of coastal food webs. This work has been and
                                    continues to be done. To preserve and conserve the resource, it is manda-
                                    tory to know how much seagrass is present and, further, to determine the
                                    rate of relative loss or gain both locally and regionally. Any "no net loss"
                                    criterion is meaningless if distribution and abundance maps are missing.

                                    ENVIRONMENTAL STUDIES

                                    Long-term management of natural systems demands an understanding of
                                    the effect of environmental fluctuations on these systems. For example, the
                                    impact-both immediate and cumulative-of coastal development on
                                    seagrass-dominated systems cannot be correctly interpreted until the
                                    physical and chemical characteristics necessary for plant growth and
                                    reproduction are fully understood. Although information is available for
                                    eelgrass, it is lacking for other species on this coast. Also, rates of change
                                    in population size and characteristics within coastal waters near areas of
                                    urban, industrial, or agricultural activity can be more easily explained
                                    when we have comprehensive knowledge of the effect of random natural
                                    disturbances on seagrass systems. Moreover, the relationship between
                                    changing environmental conditions, seagrass patch size, and animal
                                    distribution and density needs to be clarified. Finally, seagrass transplant-
                                    ing has been marginally successful along the West Coast. This is a critical
                                    issue. For example, some Superfund sites (highly contaminated sites
                                    designated for special attention and cleanup by the U.S. Environmental
                                    Protection Agency) located in estuaries and coastal areas demand restora-
                                    tion of viable seagrass systems as part of overall project goals. To comply
                                    with these goals, managers and scientists must develop the best configura-
                                    tion of preserved or created habitats. Unless transplants become more
                                    successful, this will not be possible.

                                      Coastal development and seagrass distribution and abundance
                                    Some human activities (e.g., dredging and filling and dock construction)
                                    directly impact seagrasses in this region. Also, non-point source contami-
                                    nants and suspended sediment resulting from activities connected with
                                    agriculture, logging, and industry affect the condition of soft-bottom and
                                    rocky coast habitats. The cumulative effect of these phenomena on the






                                                                                     Research Gaps and Needs 13


                                     distribution and abundance of seagrasses remains unknown. Long time-
                                     series environmental data sets are needed in a variety of seagrass environ-
                                     ments so that models describing the impacts of human activity can be
                                     generated and tested and rational management decisions can be made.

                                     - Natural disturbance and patch size
                                     The impact of stochastic events (e.g., winter storms and ice scour) on the
                                     long-term viability of seagrass meadows has not been determined. Conse-
                                     quently, we have little knowledge of long-term variation in the boundaries
                                     of meadows or patches. More information is needed from areas where the
                                     effects of human-induced disturbance is reduced. These types of data will
                                     enable us to interpret more accurately the responses of seagrasses to
                                     chronic stress.


                                     9 Patch size and animal distribution and density
                                     Efforts must be made to discern the relationship between patch or meadow
                                     size and animal distribution and density. Above-ground biomass is impor-
                                     tant not only for shelter and foraging but also as substrate for grazing and
                                     spawning. It is clear that seagrasses are an important link in the continued
                                     survival of many nearshore animals. What is not clear is the quantitative
                                     link between seagrass patch size and many animal species. Consequently,
                                     researchers are not able to state with confidence the critical patch size
                                     necessary to sustain important commercial species such as Pacific herring
                                     (Clupea harengus pallasi) or Dungeness crab (Cancer magister). This
                                     information is necessary to realistically assess the functional equivalence of
                                     restored or newly established seagrass meadows.

                                     e Seagrass transplant requirements
                                     Results from reports of failed eelgrass mitigation transplants (especially
                                     transplants greater than I hectare), particularly in the Pacific Northwest,
                                     suggest that these plants cannot be successfully transplanted (Thom 1990).
                                     This may be based on false assumptions. Many projects failed because the
                                     site was inappropriate. Commonly, eelgrass was not growing at the
                                     transplant site and no modifications were made to create conditions that
                                     would promote the growth and reproduction of transplanted stock (Thom
                                     1990).






                                       14 Managing Seagrass Systems in Western North America


                                           It is also necessary to identify appropriate transplant techniques (e.g.,
                                       bare root, anchored shoot bundles, sediment plugs containing whole
                                       plants, or seeds), Information imported from studies outlined above in the
                                       sections on Physiology (e.g., root and shoot nutrient physiology) and
                                       Population Dynamics and Reproduction (e.g., role of genetic diversity and
                                       gene flow within populations) is necessary before we can confidently
                                       advocate transplant techniques to achieve maximurn restoration efficiency.
                                           There is a clear need for transplant guidelines for all species of
                                       seagrass on the West Coast. These guidelines should be regionally specific
                                       and should suggest monitoring programs and reporting protocols. Di-
                                       rected research necessary to achieve this goal includes: (1) complete
                                       understanding of environmental conditions (e.g., sediment, nutrient, and
                                       light environments) necessary for successful transplant, (2) relative
                                       importance of genetic composition and diversity for transplants, and (3)
                                       evaluation of the most appropriate techniques for large-scale transplants.


                                       Concluding Remarks

                                       As illustrated in this report, an informal network exists between seagrass
                                       scientists and resource managers along the West Coast. In addition, we
                                       have summarized research gaps and needs identified by the scientists most
                                       intimately connected with seagrass systems in this region. The job of
                                       seagrass researchers now is to link these separate but related phenomena
                                       in an effort to promote sound seagrass management for the region as a
                                       whole.
                                           We thank all contributors for providing thoughtful, measured re-
                                       sponses toward an understanding of research gaps and needs for West
                                       Coast seagrasses. We are confident that these comments, grounded in
                                       years of observation and analysis, can lend themselves to coordinated
                                       research projects built on time-series data collection and collaborative
                                       investigations. If such is the case, the formula necessary to compute the
                                       effects of human development on the preservation and creation of healthy
                                       seagrass systems in this region might be achieved.






                                                                                           Research Gaps and Needs 15


                                        References

                                        Brayshaw, T.C. 1985. Pondweeds and bur-reeds, and their relatives, of British
                                            Columbia. British Columbia Provincial Museum Occasional Paper No. 26.
                                            Province of British Columbia. 167 pp.

                                        den Hartog, C. 1970. Seagrasses of the world. Amsterdam, North Holland. 275
                                            PP.

                                        Domning, D.P. 1977. Sirenian evolution in the North Pacific Ocean. Univ. Calif.
                                            Publ. Geological Sciences, Vol. 118. 176 pp.

                                        Fredette, TJ., M.S. Fonseca, W.J. Kenworthy, and S. Wyllie-Echeverria. 1988. An
                                            investigation of eelgrass (Zastera marina) transplanting feasibility in San
                                            Francisco Bay, California. COE Report EL-88-2. U.S. Army Engineer Water-
                                            ways Experiment Station, Vicksburg, MS. 33 pp.

                                        Harrison, P.C., and R.E. Bigley. 1982. The recent introduction of the seagrass
                                            Zosterajaponica Aschers. and Craebn. to the Pacific coast of North America.
                                            Can. J. Fish. Aquat. Sci. 39:1642-1648.

                                        Kitting, C.L., and S. Wyllie-Echeverria. 1992. Seagrasses of San Francisco Bay:
                                            Status, management and conservation needs. In: Yosemite Centennial
                                            Symposium Proceedings, National Park Service, NPS D-374, pp. 388-395.
                                            Denver Service Center.

                                        Mason, H.L. 1957. A flora of the marshes of California. University of California
                                            Press, Berkeley, CA. 897 pp.

                                        Merkel, K., and R. Hoffman (eds.). 1990. Proceedings of the California Eelgrass
                                            Symposium. Sweetwater River Press (Pacific Southwest Biological Services),
                                            National City, CA. 78 pp.

                                        Nichols, F.H., and M.M. Pamatmat. 1988. The ecology of the soft bottom benthos
                                            of San Francisco Bay: A community profile. U.S. Fish and Wildlife Service
                                            Biological Report 85(7.19). 73 pp.

                                        Phillips, R.C., and E.G. Menez. 1988. Seagrasses. Smithsonian Contributions to
                                            the Marine Sciences No. 34. 104 pp.

                                        Phillips, R.C., and S. Wyllie-Echeverria. 1990. Zostera asiatica Miki on the Pacific
                                            Coast of North America. Pacific Science 44(2):130-134.

                                        Puget Sound Water Quality Authority. 1989. Recommendations on developing a
                                            geographic information system for Puget Sound and updating the Puget
                                            Sound Atlas. Puget Sound Water Quality Authority, U.S. EPA, Region 10,
                                            Seattle. 43 pp.

                                        Setchell, W.A. 1929. Morphological and phenological notes on Zostera marina L.
                                            Univ. Calif. Publ. Bot. 14:185-201.






                                         16 Managing Seagrass Systems in Western North America


                                         Steward, AX, L.J. Dennis, and H.M. Gilkey. 1963. Aquatic plants of the Pacific
                                             Northwest. Oregon State University Press, Corvallis, OR. 261 pp.

                                         Thom, R.M. 1990. A review of eelgrass (Zostera marina L.) transplanting projects
                                             in the Pacific Northwest. The Northwest Environmental Journal 6:121-137.

                                         Wyllie-Echeverria, S., and P. J. Rutten. 1989. Inventory of eelgrass (Zostera
                                             marina L.) in San Francisco/San Pablo Bay. National Marine Fisheries
                                             Service, Southwest Region Admin. Report SWR-89-05. 18 pp.

                                         Wyllie-Echeverria, S. 1990. Geographic range and distribution of Zostera marina,
                                             eelgrass, in San Francisco Bay, California. In: K. Merkel and R. Hoffman (eds.),
                                             Proceedings of the California Eelgrass Symposium, pp. 65-69. Sweetwater
                                             River Press (Pacific Southwest Biological Services), National City, CA.

                                         Zimmerman, R.C., J.J. Reguzzoni, S. Wyllie-Echeverria, M. Josselyn, and R.S.
                                             Alberte. 1991. Assessment of environmental suitability for growth of Zostera
                                             marina L. (eelgrass) in San Francisco Bay. Aquat. Bot. 39:353-366.


                                         Appendix

                                         The following sections describe seagrass research since 1985 in Southwest-
                                         ern British Columbia, Canada, and in Baja California, Mexico.


                                         SEAGRAss RESEARCH IN SOUTHWESTERN BRITISH
                                         COLUMBIA, CANADA

                                         Paul G. Harrison
                                         Studies have focused on three levels of the ecological hierarchy: commu-
                                         nity, ecosystem, and population. Population-level studies began with basic
                                         demographic questions (e.g., Bigley and Harrison 1986), developed into
                                         applied areas (transplantation), and recently returned to basic aspects of
                                         population establishment and growth.
                                              Initially, community-level studies focused on areas where two Zostera
                                         species overlap (Zostera marina and Z. japonica). This phenomenon was
                                         due in part to port construction, which modified the physical environment
                                         and resulted in the expansion of Z. marina into a zone formerly occupied
                                         by Z. japonica. This in turn led to major changes in infaunal populations
                                         (Harrison 1987). Later, ecosystem-level studies explored the details of
                                         detritus production from Z. marina leaves and its consumption by mi-
                                         crobes and invertebrates (Harrison 1989). As an adjunct to this research,






                                                                                                Research Gaps and Needs 17


                                          some physiological studies were made on translocation and nutrient
                                          dynamics (Heminga et al. 1991).
                                               Finally, loss of Z. marina, principally caused by erosion related to port
                                          construction, led to transplant technique experiments (Harrison 1990a,
                                          1990b). These transplants were successful in that patches persisted or
                                          expanded during the monitoring period (5 years). However, several years
                                          later some transplants had perished. Consequently, recent studies have
                                          again focused on detailed demographic topics (e.g., shoot initiation and
                                          longevity, seed banks, germination) in an attempt to understand the
                                          mechanisms that control population dynamics (Nomme and Harrison
                                          1991a, 1991b; Harrison 1990c). In addition, clonal structure (Harrison and
                                          Durance 1992), recognized as a basic but poorly understood aspect of
                                          seagrass populations, will be the subject of future studies.

                                          References
                                          Bigley, R.E., and Harrison, P.G. 1986. Shoot demography and morphology of
                                              Zosterajaponica and Ruppia maritima from British Columbia, Canada,
                                              Aquat. Bot. 24:69-82.

                                          Harrison, P.G. 1987. Natural expansion and experimental manipulation of
                                              seagrass (Zostera spp.) abundance and the response of infaunal invertebrates.
                                              Estuar. Coast. Shelf Sci. 24:799-812.

                                          Harrison, P.C. 1989. Detrital processing in seagrass systems: A review of factors
                                              affecting decay rates, remineralization, and detritivory. Aquat. Bot. 35:263-
                                              288.

                                          Harrison, P.G. 1990a. Experimental eelgrass transplants in southwestern British
                                              Columbia, Canada. In: K. Merkel and R. Hoffman (eds.), Proceedings of the
                                              California Eelgrass Symposium, pp. 46-57. Sweetwater River Press (Pacific
                                              Southwest Biological Services), National City, CA.

                                          Harrison, P.G. 1990b. Variations in success of eelgrass transplants over a five-
                                              years' period. Environ. Conserv. 17:157-163.

                                          Harrison, P.G. 1990c. Population dynamics of seagrasses. In: R.C. Phillips and
                                              C.P. McRoy (eds.), Seagrass research methods. UNESCO Monographs on
                                              Oceanographic Methodology, Vol. 9, pp. 27-32.

                                          Harrison, P.G., and C. Durance. 1992. Variation in clonal structure in an eelgrass
                                              (Zostera marina) meadow on the Pacific coast of Canada. Can. J. Bot. 70:653-
                                              657.

                                          Hemminga, M.A., P.G. Harrison, and F. van Lent. 1991. The balance of nutrient
                                              losses and gains in seagrass meadows. Mar. Ecol. Prog. Ser. 71:85-96.






                                       18 Managing Seagrass Systems in Western North America


                                       Nomme, K.M., and P.C. Harrison. 1991a. A multivariate comparison of the
                                          seagrasses Zostera marina and Zostera japonica in monospecific versus mixed
                                          populations. Can. J. Bot. 69:1984-1990.

                                       Nomme, K.M., and P.G. Harrison. 1991b. Evidence for interaction between the
                                          seagrasses Zostera marina and Zosterajaponica on the Pacific coast of
                                          Canada. Can. J. Bot. 69:2004-2010.


                                       SEAGRAss RESEARCH IN BAJA CALIFORNIA, MEXICO

                                       Silvia E. Ibarra-0bando
                                       Dawson (1962) was the first to describe the presence of extensive eelgrass
                                       (Zostera marina L) meadows in Baja California, specifically in San Quintfn
                                       Bay on the Pacific Coast of Baja California. Barnard (1970) conducted
                                       detailed investigations of the invertebrates associated with eelgrass.
                                       Further descriptions are included in reports by Neuenschwander et al.
                                       (1979) and Kramer (1976). Kramer noted that maximum eelgrass biomass
                                       occurs during winter and spring, coinciding with the arrival of the black
                                       brant (Branta bernicla nigricans) from its summer feeding and breeding
                                       grounds in the Arctic. Eelgrass is an important food for wintering brant at
                                       several sites in Baja California (Kramer 1976, Ward 1983).
                                           On the Pacific side of Baja California, Zostera marina extends south as
                                       far as Bahia Magdalena (240N) (Saunders and Saunders 1981). In these
                                       temperate waters, eelgrass is perennial (Harrison 1982); however, in the
                                       warm waters of the Gulf of California, an annual variety grows (Phillips
                                       and Backman 1983). Descriptive and functional information on eelgrass
                                       beds in Baja California is currently only fragmentary.
                                           With limited funding, my laboratory in 1982 began a complete
                                       descriptive study of the seagrasses in Baja California. For several years, we
                                       collected data for a number of plant characteristics (e.g., above and
                                       underground biomass, blade production, and shoot dynamics) and envi-
                                       ronmental parameters (e.g., tidal elevation and sediment characteristics) as
                                       well as epiphyte identification and biomass. These findings, summarized in
                                       Ibarra-Obando (1989), allow us to theorize that eelgrass prairies in San
                                       Quintfn Bay are highly dynamic communities with a distinctive seasonal
                                       cycle that appears to differ from other eelgrass systems at more northerly
                                       locations along the West Coast of North America.






                                                                                             Research Gaps and Needs 19


                                              Current efforts focus on the role seagrass prairies play as nursery
                                         grounds for juvenile fish and invertebrates. To date, no studies have
                                         described the relationship between fish and eelgrass beds in Baja
                                         California.
                                             Based on past and current studies, the following research gaps have
                                         been identified for seagrass systems in Baja California, Mexico: (1) charac-
                                         terization of the relationship between eelgrass and macro- and microalgae
                                         with respect to light and space; (2) determination of the relative contribu-
                                         tions of eelgrass, salt marsh systems, and macro- and microalgae (phy-
                                         toplankton included) to organic matter production; (3) determination of
                                         the relative influence of various sediment regimes (e.g., sand, silt) on plant
                                         morphology; (4) comparison of the structure and function of annual versus
                                         perennial eelgrass meadows; (5) descriptions of root and rhizome morphol-
                                         ogy across tidal elevations and determination of the relative functions of
                                         the various morphs; (6) detailed descriptions of the reproductive cycle in
                                         Baja waters; and (7) initiation of comprehensive research projects, similar
                                         to the studies in San Quintfn Bay, at more southerly locations.
                                              Compared to other locations along the Pacific Coast of North America,
                                         the Baja California coast is still relatively undisturbed. Scientific interest in
                                         this geographic region turns on the fact that undisturbed communities can
                                         provide basic information on ecosystem structure and functioning. These
                                         data are useful for regional seagrass management. Also, international
                                         efforts to study and preserve eelgrass beds in Mexico are needed. Establish-
                                         ing binational research projects would not only accomplish this goal but
                                         also provide the kind of information needed to meet existing knowledge
                                         gaps.

                                         References
                                         Barnard, J.L. 1970. Benthic ecology of Bahia de San Quintfn, Baja California.
                                             Smithsonian Contributions to Zoology, 44. Smithsonian Institution Press,
                                             Washington. 60 pp.

                                         Dawson, E.Y. 1962. Benthic marine exploration of Bahia de San Quintfn, Baja
                                             California, 1960-61. No. 7. Marine and marsh vegetation. Pac. Natl. 3:275-280.

                                         Harrison, P.C. 1982. Spatial and temporal patterns in abundance of two intertidal
                                             seagrasses, Zostera americana Den Hartog and Zostera marina L. Aquat. Bot.
                                             12:305-320.






                                        20 Managing Seagrass Systems in Western North America


                                        lbarra-Obando, S.E. 1989. Las praderas de pastos marinos del Pacffico mexicano
                                           con 6nfasis en Zostera marina Linneo. Cap. 1.:1-20. In: Rosa-V61ez, J. de la y
                                           F. Gonzdlez-Farias (eds.), T@mas de Oceanograffa Biol6gica en Mexico.
                                           Universidad Aut6noma de Baja California. Ensenada. 337 pp.

                                        Kramer, G.W. 1976. Winter ecology of black brant at San Quintfn Bay, Baja
                                           California, Mexico. M.S. thesis, Humboldt State University, Arcata, CA. 80 pp.

                                        Neuenschwander, L.F., T.H. Thorsted, Jr., and R.J. Vogl. 1979. The salt marsh and
                                           transitional vegetation of Bahia de San Quintfn. Bull. Southern California
                                           Acad. Sci. 78:163-182.

                                        Phillips, R.C., and T.W. Backman. 1983. Phenology and reproductive biology of
                                           eelgrass (Zostera marina L.) at Bahia Kino, Sea of Cortez, Mexico. Aquat. Bot.
                                           17:85-90.

                                        Saunders, G.B., and D.C. Saunders. 1981. Waterfowl and their wintering grounds
                                           in Mexico, 1937-64. U.S. Fish and Wildlife Service Res. Publ. 138. Washing-
                                           ton, D.C. 151 pp.

                                        Ward, D.J. 1983. The relationship of two seagrasses: Zostera marina and Ruppia
                                           maritima to black brant, Branta bernicla nigricans, in San Ignacio Lagoon,
                                           Baja California, Mexico. M.S. thesis, University of Oregon, Eugene. 57 pp.






                                                                                                        Research Caps and Needs 21


                                              Contributors

                                              Josef Daniel Ackerman                          (U.S. mailing address for lbarra-Obando:
                                              Mechanical Engineering                         P.O. Box 434844
                                              University of Toronto                          San Diego, CA 92143-4844)
                                              5 King's College Rd.                           Christopher L. Kitting
                                              Toronto, Ontario M5S IA4
                                              Canada                                         Dept. of Biological Sciences
                                                                                             Marsh Research Station
                                              Randall S. Alberte                             California State University
                                              Office of Naval Research                       Hayward, CA 94542
                                              800 N. Quincy St.
                                              Arlington, VA 22217-5000                       Keith W. Merkel
                                                                                             Pacific Southwest Biological Services
                                              Dan Baron                                      P.O. Box 985
                                              Biological Sciences Dept.                      National City, CA 92050
                                              California State University
                                              Hayward, CA 94542                              Ronald C. Phillips
                                                                                             Battelle-Pacific Northwest Laboratory
                                              Douglas Bulthuis                               P.O. Box 999; MSIN P7-68
                                              Padilla Bay Natl. Estuarine Research           Richland, WA 99352
                                                Reserve
                                              1043 Bay View-Edison Rd.                       Mary Ruckleshouse
                                              Mt. Vernon, WA 98273                           Dept, of Botany KB-15
                                                                                             University of Washington
                                              Paul A. Cox                                    Seattle, WA 98195
                                              Dept. of Botany and Range Science
                                              Brigham Young University                       Charles A. Simenstad
                                              Provo, UT 84602                                Wetland Ecosystem Team
                                                                                             School of Fisheries VM-10
                                              LoAnn Hallurn                                  University of Washington
                                              Wetland Ecosystem Team                         Seattle, WA 98195
                                              School of Fisheries WH-10
                                              University of Washington                       Ronald M. Thom
                                              Seattle, WA 98195                              Battelle/Marine Sciences Laboratory
                                                                                             1529 W. Sequim Bay Road
                                              Paul G. Harrison                               Sequim, WA 98382
                                              Dept. of Botany
                                              University of British Columbia                 Susan L. Williams
                                              Vancouver, B.C. V6T 1Z4                        Biology Dept.
                                              Canada                                         San Diego State University
                                                                                             San Diego, CA 92182-0057
                                              Robert F. Hoffman
                                              Natl. Marine Fisheries Service, SW Reg.        Sandy Wyllie-Echeverria
                                              501 West Ocean Boulevard, Suite 4200           School of Marine Affairs HF-05
                                              Long Beach, CA 90802-4213                      University of Washington
                                                                                             Seattle, WA 98195
                                              Silvia E. lbarra-Obando
                                              Centro de Investigaci6n Cientifica y           Richard C. Zimmerman
                                                Educaci6n Superior de Ensenada               Hopkins Marine Station
                                              Apdo. Postal 2732                              Pacific Grove, CA 93950
                                              Ensenada, B.C.
                                              Mexico



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