Maintaining Ecological Integrity of our Parks • Maintenir l’intégrité écologique de nos parcs

Parks Canada-Technical Reports in Ecosystem ScienceParcs Canada-Rapports techniques en matière de sciences des écosystèmes

Living Marine Legacy of Gwaii Haanas. I:Marine Plant Baseline to 1999 and Plant-related Management Issues

N. A. Sloan and P. M. Bartier

March 2000Report 027

Atlantic Region, Parks Canada is producing three reportseries in ecosystem science. They are intended tocommunicate new scientific information, documentscientific data, summarize existing knowledge, or offertechnical recommendations. The primary function and theintended audience of a report determine the series inwhich it will be published. Each report series containsscientific and technical information that contributes toexisting knowledge but is not in a form suitable for theprimary journal literature.

• Parks Canada-Technical Reports in EcosystemScience promote a wide distribution of scientific andtechnical information from Parks Canada’secosystem investigations. The subject matter, and theseries reflect the broad interest and policies of ParksCanada in ecosystem science. This series includessuch work as, ecosystem and resource inventories,studies, surveys of species or guilds, and innovativemanagement concepts.

• Parks Canada - Ecosystem Science Review Reportsprovides a forum for literature reviews,bibliographies, and reviews of management optionsthat are often prepared for or by Parks Canada.Resource Descriptions and Analyses or chaptersthereof will be published in this series.

• Parks Canada - Ecosystem Monitoring and DataReports provide a medium for filing and achievingdata compilations where little or no analysis isincluded. Such compilations commonly are preparedin support of primary publications or TechnicalReports in Ecosystem Science. Raw data notavailable in a national data base and considered wortharchiving is published as a Parks Canada - EcosystemMonitoring and Data Report.

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Neil MunroReport Series EditorParks CanadaHistoric PropertiesHalifax, Nova ScotiaB3J 1S9(902) 426-2797FAX 426-2728

Parks Canada

Technical Reports in Ecosystem Science

Report # 27

Living Marine Legacy of Gwaii Haanas I: Marine Plant Baseline to 1999

and Plant-related Management Issues

by

N.A. Sloan and P.M. Bartier

2000

Ecosystem Management SectionGwaii Haanas National Park Reserve / Haida Heritage Site

P.O. Box 37, Queen Charlotte, BC CANADA V0T 1S0

© Her Majesty the Queen in Right of Canada,represented by the Chief Executive Officer of Parks Canada, 2000.

Canadian Cataloguing in Publication Data

Sloan, N.A.

Living marine legacy of Gwaii Haanas I :marine plant baseline to 1999 and plant-related management issues

(Parks Canada - Technical Reports in Ecosystem Science,ISSN 1200-3298 ; no. 27)

Includes an abstract in French.Includes bibliographical references.

ISBN 0-662-28529-8Cat. no. R61-2/19-27-2000E

1. Marine plants — British Columbia — Queen Charlotte Islands.2. Marine plants — British Columbia —

Gwaii Haanas National Park Reserve.3. Biotic communities — British Columbia — Queen Charlotte Islands.4. Biotic communities — British Columbia —

Gwaii Haanas National Park Reserve.5. Gwaii Haanas National Park Reserve (B.C.) — Management.

I. Bartier, P. (Patrick), 1960- .II. Parks Canada. Atlantic Region.

III. Title.IV. Series: Technical reports in ecoystem science; no. 27.

QK938.C6S56 2000 579’.177 0-980029-8

ii

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ABSTRACT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vRESUME . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vACKNOWLEDGEMENTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viPREFACE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viiEXECUTIVE SUMMARY . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . viiiINTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 SEAWEEDS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 SEAGRASSES . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 MARINE LICHENS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 HAIDA MARINE PLANT USE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6RESULTS AND DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 SPECIES LISTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 Seagrasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 Marine Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 GEOGRAPHIC DISTRIBUTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Algae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 Seagrasses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 MARINE PLANT ISSUES IN GWAII HAANAS MANAGEMENT . . . . . . . . . . . . . . 19 Nearshore Marine Environmental Monitoring . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20 Intertidal Seaweed and Subtidal Kelp Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21 Seagrass Meadows . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Marine Lichens . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25 Kelp Forest Ecosystem Issues . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Sea Otters and Haida Gwaii History . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26 Kelp Deforestation, Sea Otter and Red Sea Urchin . . . . . . . . . . . . . . . . . . . . . . . . . 26 Kelp Forest-associated Fisheries . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 28 Pacific Herring Spawn-on-Kelp Fisheries and Kelp Mariculture . . . . . . . . . . . . . . . . 29 Visitor Impacts – Trampling Intertidal Seaweeds . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30CONCLUSIONS and RECOMMENDATIONS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31 GWAII HAANAS MARINE FLORA KNOWLEDGE . . . . . . . . . . . . . . . . . . . . . . . . . 31 GWAII HAANAS ECOSYSTEM MANAGEMENT . . . . . . . . . . . . . . . . . . . . . . . . . . 32LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33

Figure 1. Location of Gwaii Haanas within Haida Gwaii (Queen Charlotte Islands) includingplace names mentioned in the text. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Figure 2. The 107 shore observation locations from which marine plant data were recorded inthe coastal biophysical inventory of Gwaii Haanas by Harper et al. (1994). . . . . . . . 7

Figure 3. Sample locations from surveys of Gwaii Haanas commissioned by EnvironmentCanada – Parks. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

Figure 4. Sample locations in Haida Gwaii from which marine plant specimens are retained atthe University of British Columbia Phycological Herbarium. . . . . . . . . . . . . . . . . . . 8

Figure 5. Sample locations of various marine surveys in Haida Gwaii from which marineplants were identified. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

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Figure 6. The distribution of kelp symbols (likely representing mostly Macrocystisintegrifolia and Nereocystis luetkeana) on six Canadian Hydrographic Service nauticalcharts of the Moresby Island area. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Figure 7. Inferred distribution of kelp (likely dominated by Macrocystis integrifolia andNereocystis luetkeana) from the coastal biophysical inventory of Harper et al.(1994). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12

Figure 8. Locations of the 19 wave exposure assessment sites in the SGaang Gwaiiarchipelago surveyed in July, 1999 by Druehl and Hopkins (1999). . . . . . . . . . . . . . 15

Figure 9. Locations of the six permanent intertidal seaweed monitoring transects andMacrocystis sp. observations made in July, 1999 by Druehl and Hopkins (1999).

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Figure 10. Distribution of Sargassum muticum from various marine surveys in Haida Gwaii.17Figure 11. Distribution of eelgrass (Zostera marina) and surfgrasses (Phyllospadix spp.) from

various surveys in Haida Gwaii. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18Figure 12. An example of eelgrass (Zostera marina) mapping in Louscoone Inlet, summer,

1999. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

Table 1. Haida names for some marine plants from Haida Gwaii. . . . . . . . . . . . . . . . . . . . . 5Table 2. Herbarium marine specimens from Haida Gwaii in the Canadian Museum of Nature,

Ottawa. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10Table 3. Seaweed and seagrass taxa of Haida Gwaii and the Oregon to Southeast Alaska

region. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Table 4. Intertidal algae, seagrass and lichens, used with exposure, substrate and tidal zone,

to discriminate community types from aerial photographs. . . . . . . . . . . . . . . . . . . . 14Table 5. The expected presence/absence of marine plant species indicating wave exposure on

SGaang Gwaii. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16Table 6. The number (and percent) of sites at which seagrasses occurred according to

exposure category at 254 sites in Gwaii Haanas. . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Table 7. Summary of selected intertidal monitoring attributes involving marine plants in

National Parks and Monuments from Alaska to California. . . . . . . . . . . . . . . . . . . . 23

Appendix A. Herbaria staff contacted in the preparation of this report. . . . . . . . . . . . . . . . . 45Appendix B Part 1. Number of sites at which all marine algae and seagrasses have been

reported from Haida Gwaii and Gwaii Haanas National Park Reserve. . . . . . . . . . . 46Appendix B Part 2. Number of specimens of marine lichens according to shore zones and

substrates in Haida Gwaii. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57Appendix C. Distribution maps of individual algal and seagrass taxa in Haida Gwaii.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59

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Sloan, N.A. and P. M. Bartier, 2000. Living Marine Legacy of Gwaii Haanas I: Marine Plant Baselineto 1999 and Plant-related Management Issues. Pks. Can. - Tech. Rep. Eco. Sci., No. 27, viii + 104 p.

ABSTRACT

This is the first report in a series of baseline marine biological inventories for the Haida Gwaii(Queen Charlotte Islands) archipelago including Gwaii Haanas National Park Reserve/HaidaHeritage Site. We list the seaweed and seagrass species and map their distributions known tothe end of 1999. Our geographic information system contains records of 348 seaweed andfour seagrass species from 456 localities in the archipelago. We include a preliminary marinelichen flora of 88 species. The importance of plant communities to the coastal ecosystemintegrity of Gwaii Haanas is great. Kelp (large seaweed) forests and seagrass meadowscomprise ecosystems that shelter many species of juvenile and adult fishes and invertebrates. Kelp is critical to traditional and commercial Pacific herring spawn-on-kelp fisheries. Further,marine plants are important to indigenous Haida culture.

RESUMÈ

Il s’agit du premier d’une série de rapports de base sur l’inventaire des espèces marines del’archipel Haida Gwaii (îles de la Reine-Charlotte), qui comprend la réserve de parc national/lesite du patrimoine haïda Gwaii Haanas. Nous y énumérons les diverses espèces d’algues et degraminées marines et y cartographions leurs aires de distribution connues jusqu’à la fin de1999. Notre système d’information géographique contient une liste de 348 espèces d’algues etde 4 espèces de graminées marines peuplant 456 localités de l’archipel. Le rapport renfermeégalement une liste préliminaire de 88 espèces de lichens marins. Les communautés végétalescontribuent grandement à l’intégrité de l’écosystème côtier de Gwaii Haanas. Les forêts devarech (grandes algues) et les prés de graminées marines forment des écosystèmes qui serventd’habitat à de nombreuses espèces de poissons et d’invertébrés juvéniles et adultes. Le varechjoue aussi un rôle essentiel dans la récolte traditionnelle et la pêche commerciale des œufs dehareng du Pacifique. Enfin, les plantes marines revêtent de l’importance dans la cultureindigène des Haïdas.

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ACKNOWLEDGEMENTS

Many people helped us to prepare this report. Special thanks to Dr. Louis Druehl (CanadianKelp Resources Ltd., Bamfield) for his mentoring, partnership in the field and review of thetext. An adequate species listing would not have been possible without the assistance of Dr.Michael Hawkes (Curator of the Phycological Herbarium, University of British Columbia)who reviewed the text and taxonomy. Dr. Irwin Brodo (Canadian Museum of Nature, Ottawa[CMN]) very generously provided the marine lichen list, advice on marine aerosol pollutionmonitoring and reviewed the lichen text. Dr. Robert F. Scagel (Professor Emeritus, UBC)kindly wrote the preface to provide some historical context to this project and commented onthe text. Dr. Tom Tomascik (Senior Marine Advisor, Parks Canada) kindly reviewed the text. We thank Dr. Michael Hawkes and Mr. Mike Shchepanek (CMN) for providing criticalherbarium records of their Haida Gwaii material. Dr. Paul G. Harrison (UBC) kindlyidentified a seagrass sample and provided guidance on local seagrass taxonomic problems. Mr. Russ Jones (Haida Fisheries Program) kindly advised us on Haida Nation kelp-associatedactivities. Ms. Barb Wilson (Cultural Resource Manager, Gwaii Haanas) reviewed the Haidaknowledge information. Thanks to Mr. Scott Giroux and Ms. Debbie Gardiner of GwaiiHaanas Warden Services for the eelgrass map and to Ms. Maggie Stronge for the Sargassummuticum observation. Ms. Heather Holmes (Pacific Rim National Park Reserve, Ucluelet,BC) shared information on their intertidal monitoring. Drs. Colin Levings (Department ofFisheries and Oceans, Vancouver, BC) and Ron Thom (Battelle Northwest Marine Lab,Sequim, WA) advised us on seagrass monitoring in the Pacific Northwest. We receivedinformation on nearshore marine monitoring from Mr. Gary Davis (Channel Islands NationalPark, Ventura, CA), Mr. Ed Bowlby (Olympic Coast National Marine Sanctuary, PortAngeles, WA), Dr. Megan Dethier (Friday Harbor Laboratory, Friday Harbor, WA) and Ms.Gail Irvine (Alaska Biological Science Center, Anchorage, AK). Ms. Christine Boesveld(Parks Canada, Hull) kindly helped with our print information needs. We are pleased toacknowledge the support of Mr. Brian Reader, Ecosystem Management Coordinator, GwaiiHaanas.

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PREFACE

My first exposure to the marine flora of British Columbia was in 1946 as an undergraduate atthe University of British Columbia (UBC). In the late 1940s and early 1950s I studied andcollected marine plants at many sites between the southern end of Vancouver Island andLangara Island in Haida Gwaii (Queen Charlotte Islands). I visited the extensive kelp forestssheltered in Cumshewa Inlet on the east coast of Moresby Island and saw the rugged andexposed coast at Tasu Sound off the west coast of Moresby Island and Hippa Island off thewest coast of Graham Island looking (unsuccessfully) for Postelsia palmeaformis. I alsovisited areas along the north shore of Graham Island between the Mazarredo Islands andChanal Reef off Langara Island. My most memorable Haida Gwaii experience was on thenorth coast of Graham Island, where I first observed the extent of the tidal amplitude and thegreat richness of the intertidal flora. By myself in a 3-m dinghy, observing, photographing andcollecting marine algae in Hazardous Cove on Langara Island was an especially exciting andrewarding experience. These early visits, together with those of my graduate students andpostdoctoral fellows after I joined UBC in 1952, provided opportunities to collect marinealgae for UBC’s Phycological Herbarium. Except for the somewhat questionable (as tolocale) records for a few marine algae (including Egregia menziesii, which may have beencollected somewhere in Haida Gwaii) recorded by Archibald Menzies in 1787-1788, ourcollections in the 1940s, 1950s and 1960s provided the first significant records of the benthicmarine algae of Haida Gwaii. The UBC Phycological Herbarium, which has grown from ahandful of specimens in 1946 to over 84,000 in 1999, has provided a rich resource ofphycological data for this report.

The special value of this publication is that it brings together for the first time for the area adetailed inventory of the marine flora based on published literature as well as on availableherbarium specimens. Much more needs to be done to obtain a more comprehensiveinventory of the biota of the area. A great diversity of habitats exists in Haida Gwaii andmany more sites need to be examined intertidally as well as subtidally – especially wherediving has been little exploited.

Marine plants can be useful indicators of the physio-chemical characteristics of theenvironment in which they occur. Haida Gwaii occupies a unique and important geographicposition as it presents a transition from the southern region to the northern region andcontributes significantly to an understanding of the distribution patterns of benthic marineplants on the Pacific Coast of North America. With increasing need for conservation andconcern for the marine habitat, it is especially important that we have analyses such aspresented in this publication, so that, over time, it will be possible to detect changes in marinebiota, assess the impact of man’s use and exploitation of the marine habitat, and attempt toconserve the biodiversity of this phycologically rich area.

Robert F. ScagelProfessor Emeritus, Department of Botany

University of British Columbia

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EXECUTIVE SUMMARY

“Gosh, there’s an awful lot of work to make even the most cursory survey of this sort.”Ed Ricketts on marine biological surveys; Masset Inlet, June 1946 (Hedgpeth 1978)

This is the first report in a series providing baseline marine biological inventories for the HaidaGwaii (Queen Charlotte Islands) archipelago including Gwaii Haanas National ParkReserve/Haida Heritage Site. An assessment of biodiversity is central when addressing ParksCanada’s mandate to protect and conserve representative samples of Canada’s marine areasby maintaining ecosystem structure and function.

We list the seaweed and seagrass species and map their distributions known to the end of1999. Our geographic information system (GIS) database contains 6031 records, comprising348 seaweed and four seagrass species recorded from 456 localities among the islands. HaidaGwaii is of marine biogeographical interest as it represents the southern extreme of thenorthern seaweed flora. The closest area whose flora is reasonably well known is Bamfield onthe southwestern coast of Vancouver Island. A preliminary marine lichen flora of 88 speciesis also included.

The importance of plant-structured communities to the nearshore ecosystems of Gwaii Haanascannot be overstressed. Besides being of academic service to marine botanists andbiodiversity specialists, this Haida Gwaii marine plant survey has practical implications forParks Canada’s long-term stewardship of Gwaii Haanas in particular.

Kelp forests and seagrass meadows comprise vital nearshore marine ecosystems here. Theyprovide detritus into coastal food webs, shelter many species as nurseries for juveniles andhabitat for adults. Kelp is fundamental to traditional and commercial herring spawn-on-kelpfisheries. Further, marine plants are important to Haida culture.

Issues relating to Parks Canada’s marine ecosystem mandate in Gwaii Haanas include:

• understanding the role of plants in structuring nearshore marine communities;• environmental monitoring of threats to, and well-being of, ecosystems;• assessing impacts of repatriated sea otters on nearshore kelp forest ecosystems; • evaluating the role of red sea urchin grazing (herbivory) on kelp forest ecosystems; • rehabilitation of depleted kelp-associated northern abalone stocks;• maintaining healthy kelp resources in support of herring spawn-on-kelp harvest;• appreciating the role of seagrass meadows in the land-sea linkages in estuaries; and• backcountry monitoring of visitor impacts to intertidal habitats.

Marine plant-associated management issues are discussed and recommendations madeconcerning maintaining nearshore ecosystem structure and function of Gwaii Haanas. Wehope that this report will serve regional marine botany and help focus attention on marineecosystem management issues of Gwaii Haanas.

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INTRODUCTION

Parks Canada is mandated to protect andconserve representative samples of marineregions that includes maintaining ecosystemstructure and function while permittingmultiple uses within proposed MarineConservation Areas. An important step inassessing biodiversity is creatingtaxonomically and systematically (i.e.,named and ordered) up-to-date inventories.This is the first report in a series providingbaseline marine inventories of Haida Gwaii(Queen Charlotte Islands) and particularlythe living marine legacy of Gwaii Haanas.Further, this follows up on our Marine PlantBiodiversity Plan (Druehl 1999 a).

This marine plant report addresses GwaiiHaanas’ management objectives for twomain reasons:

• to advise management on the knowledgeof current marine plant species and plant-associated ecosystem management issues;and

• to establish the marine plant historicbaseline in the form of a reliable specieslisting for specialists to augment andcompare with other North Pacific regions.

Gwaii Haanas comprises the southern end ofMoresby Island and associated islands insouthern Haida Gwaii archipelago off thenorthern British Columbia mainland coast(Figure 1). Gwaii Haanas incorporates~1,470 km2 of land, ~3,400 km2 of proposedsea space and ~1,700 km of shoreline.

Gwaii Haanas represents the NationalMarine Conservation Area Natural Regionsof Queen Charlotte Islands Shelf to the west,Hecate Strait to the east and borders theQueen Charlotte Sound region to the south

(Mercier and Mondor 1995). Gwaii Haanas’west coast is highly exposed to the openNortheast Pacific and has a narrow shelf andcontinental slope descending rapidly to>2000 m depth within 20 km offshore. Theeast coast faces Hecate Strait, which extends~70 km to the northern British Columbiamainland, and is mostly shallower than 150m.

The environment of Gwaii Haanas marinearea is temperate coniferous forests onmountainous terrain along an extremelycomplex, mostly rocky (~75%) shoreline,much of which is exposed to high winds andheavy rains in the winter storm season. Lessthan 10% of the shoreline is sandy, of whichestuarine flats account for ~6% of the totalshoreline. The entire shoreline has receivedpreliminary biophysical classification(Harper et al. 1994) and is on a geographicinformation system (GIS) platform for ourmarine knowledge.

This marine plant inventory provides amodel for subsequent GIS-based marineinventories proposed for Gwaii Haanas.These inventories, together with thecomplete science bibliography of HaidaGwaii under preparation, will provide astarting point of natural science support forfuture ecosystem-based management ofGwaii Haanas.

A sound inventory is an important steptowards understanding the ecosystem role ofmarine plants in nearshore Gwaii Haanashabitats. At present, this role is virtuallyunknown and, arguably, would remainunknowable if such basic biodiversityinformation is not made available.

This report focuses on seaweeds, seagrassesand marine lichens only. Beach grasses,herbs and sedges rooted in sandy

2

CanadaAK

WA

OR

CA

BritishColumbia

GrahamIsland

MoresbyIsland

Gwaii HaanasNational Park Reserve/Haida Heritage Site

RennellSound

Masset

QueenCharlotte

City

LangaraIsland

SGaangGwaii

52 N 52 N

53 N 53 N

54 N

133 W 132 W 131 W

132 W

Hecate

Strait

MassetInlet

Tasu Sound

CumshewaInlet

BurnabyNarrows

Murchison-FaradayPassage

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Figure 1. Location of Gwaii Haanas within Haida Gwaii (Queen Charlotte Islands) including place names mentioned in the text.

3

backshores, estuarine wetlands flora andphytoplankton are not included, but musteventually be covered for a complete GwaiiHaanas marine plant baseline. Surveys ofbeach and wetland plant communities areavailable in specific Haida Gwaii reportssuch as Calder and Taylor (1968; MaritimeCommunities section, p. 55-71) and Ogilvie(1994); as well as general B.C. coastal floralsummaries (Klinka et al. 1989; Pojar andMacKinnon 1994; Turner 1995).

Highest quality, standardised listingsacceptable to international convention arevital to the long-term utility of any speciessurvey. Our commitment to conventionsand protocols concerning nomenclature andsystematics are described in the Methodssection.

The importance of plant-structuredcommunities to the nearshore ecosystemintegrity of Gwaii Haanas is fundamental.Kelp (large brown seaweed) forests andseagrass meadows comprise vital nearshoremarine ecosystems. Besides being ofacademic service to marine botanists andbiodiversity specialists, this report haspractical implications for our long-termstewardship of Gwaii Haanas. Further,marine plants are important to indigenous(Haida) culture.

This is the first comprehensive marine plantsurvey of this type for a national park inNorth America. In a survey of 252 NationalParks and National Monuments under U.S.National Park Service management,Stohlgren et al. (1994) reported that fewparks had complete inventories oforganisms. Further, they concluded thatinvertebrate and non-vascular plantinventories were “poor or non-existent”.

SEAWEEDS

“Seaweeds” in this report are algae; plantsthat have always lived in water. They haveno vascular system, roots, flowers or seeds.All species are characterized by alternatinglife history phases which can be isomorphicor heteromorphic. Seaweeds occurwherever there is stable attachment surface(including other organisms) from theimmediate spray zone just above the upperintertidal of rocky shores down to ~35 mbelow the zero tide line in the NortheastPacific. The lower limits of seaweeds areoften dictated by suitable substrate ratherthan amount of available light. There arethree main types of seaweed: green, red andbrown.

British Columbia’s general seaweedhandbook is Scagel (1971) and for SoutheastAlaska is O’Clair and Lindstrom (2000) .Hawkes at al. (1978) is the only broadnorthern British Columbia seaweedsynopsis. There are only a few additions tothe peer-reviewed Haida Gwaii literature,such as the Langara Island survey ofGarbary et al. (1980). Historical inventoriesof commercial British Columbia seaweedshave focused on south coast kelp (Cameron1916; Scagel 1947). The only publishedcommercial survey in Haida Gwaii wasrestricted to the north and west coasts ofGraham Island (Coon et al. 1979).

Kelp is ubiquitous in the archipelago. TheHaida have harvested k’aaw, Pacific herring(Clupea harengus pallasi) spawn-on-kelp inseason, likely for millennia, particularlyalong the east and north coasts of HaidaGwaii. In the signing of the South MoresbyAgreement between Canada and theProvince of British Columbia in July 1988,the intent to transfer the stewardship of allprovincial marine resources in Gwaii Haanas

4

to Parks Canada was declared. The transferwill occur after passage of the proposedMarine Conservation Areas Act expected in2000. Resources covered in the transfer willinclude licensing giant kelp (Macrocytsisintegrifolia) harvest in support of thecommercial J-licence, spawn-on-kelp (SOK)fishery that currently has 46 licencesProvince-wide.

SEAGRASSES

Seagrasses originate from land plants andthey flower, pollinate, produce fruit anddisperse seeds as do aquatic land grasses.They are not closely related to land grassessuch as those seen on sand/gravel beachbackshores. Unlike seaweeds, seagrasseshave true roots, leaves and an internaltubular transport (vascular) system fornutrients and gasses. In tide pools, lowerintertidal and shallow subtidal sandy tomuddy shores they can form productivemeadows that provide vital animal habitatand organic detritus primarily into inshorefood webs. Seagrasses spread vegetativelyby horizontal underground stems (rhizomes)which, with the roots, form dense mats inthe substrate. Broad-scale dispersal occursby sexual reproduction involving seed-release from pollinated flowers.

Seagrasses root in moderately to verysheltered sediment shores, although somespecies prefer sediments in crevices ofexposed rocky shores. Seagrasses occur inthe lower intertidal, down to a maximum inGwaii Haanas waters of ~10 m depth. Theupper limit of seagrasses, which is in thelower intertidal, is imposed by exposure todesiccation and wave energy. Their lower(subtidal) limit is imposed by the penetrationof sufficient light to allow photosynthesis toexceed respiration.

Seagrass meadows are found along thewave-exposed rocky British Columbia (andGwaii Haanas) coasts to sheltered,depositional areas such as bays and deltas atthe heads of inlets. Although sedimentaryhabitats are relatively small overallcompared to rocky habitats, they areconsidered very important for juvenile fishsuch as salmon, many invertebrates andmarine birds.

MARINE LICHENS

Lichens are a symbiotic partnership betweena fungus and a green alga or, lesscommonly, a cyanobacterium (previouslytermed “blue-green” algae). A brown algalsymbiot in the marine lichen, Verrucariatavaerisae, was recently reported fromCalifornia and red algal-fungal obligateassociations called mycophycobioses arealso known (M. Hawkes, personalcommunication). The fungus supplies thestructural support in lichens and the alga isthe photosynthetic component; usuallysandwiched as a distinct layer within fungallayers. Marine species grow mostly in twoof the three main growth forms; crustose andfoliose (O’Clair et al. 1996).

According to O’Clair et al. (1996), marinelichens in the Northeast Pacific are hardy,salt-tolerant and usually slow-growingspecies which firmly attach to hardsubstrates such as rock, driftwood and treebark from the upper intertidal to the forestedge. There are a number of zonationschemes for shore lichens and these must betempered by local conditions (Ryan 1988,Figure 1). For example, local freshwaterseepage can obscure marine-controlledzonation patterns (Ryan 1988). The zonesfrom Southeast Alaska are defined byO’Clair et al. (1996) as follows:

5

• “salt spray” zone - areas experiencingonly seawater spray;

• “splash” zone - areas covered by only thehighest annual tides and/or wetted byintermittent wave splash; and

• “high intertidal” zone - areas regularlycovered by seawater during high tides.

The first zone comprises the generalmaritime flora with the splash zoneseparating the maritime from the true marine(most salt tolerant) flora. Knowledge ofmarine lichens in Haida Gwaii comes almostentirely from Dr. I.M. Brodo of theCanadian Museum of Nature (e.g., Brodo1995; Brodo and Santesson 1997).Appropriate zonation of marine lichens inGwaii Haanas awaits field verification byDr. Brodo scheduled for summer, 2000.

HAIDA MARINE PLANT USE

The importance of marine plants to Haidaculture is clear when you consider that thename of T’aanuu village means eelgrass –referring to eelgrass meadows growingnearby.

The Haida Fisheries Program has gathered(and partially mapped) giant kelp (M.integrifolia) data from the east and northcoasts of Haida Gwaii in support of thetraditional spawn-on-kelp fishery (RussJones, personal communication). As well,the Haida Fisheries Program has alsocollected kelp information under contract tothe commercial J-Licence Pacific herringspawn-on-kelp fishers.

Listed in Table 1 are marine plant speciesmatched with Haida names currently in thescientific literature. This would likely be

1

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the bull kelp stipe is called (“fishing line”) (Turner 1974)other “kelps” in this general taxonomic category include species of

and as well as blades, but is itself giant kelp (Turner 1974)the species most commonly used (Turner 1995) the seaweeds gathered in early spring travels from winter villages in search of greens and sea

vegetables or “winter” seaweeds (Barb Wilson, )

tle’gaay

ngaalLaminaria, Pleurophycus,

Costaria, Egregia, Alaria Agarum Nereocystis

Porphyra

personal communication

Table 1. Haida names for some marine plants from Haida Gwaii.

Haida Name Common Name Scientific NameIhkyaa’maa1 Bull kelp Nereocystis luetkeana

Ngaal2 Giant kelp Macrocystis integrifoliaT’al Rockweed / Sea wrack Fucus gardneri

Sgyuu Red laver / Nori Porphyra perforataSgyuu Black laver / Black seaweed Porphyra abbottae3

Sgiiwaay “Winter seaweed”4 Porphyra lanceolataPorphyra torta

Ga7aan Ribbon kelp / Wing kelp Alaria marginataSgii’naaw Sea lettuce / Sea hair Ulva lactuca

Enteromorpha intestinalisChaagaan-xiilaay Coralline and some other red algae Corallina spp.

Constantinea spp.T’aask’aat’uugaa Dead man’s fingers Halosaccion glandiforme

T’aanuu Eelgrass / Seagrass Zostera marinaT’aanuu Surfgrass / Seagrass Phyllospadix spp.

6

expanded considerably with a focusedtraditional knowledge survey. Among theplants, the most important food species aregiant kelp, black laver and winter seaweed.Seaweed (sgyuu) and spawn-on-kelp(k’aaw) [using giant kelp ngaal], besidesbeing important foods to be consumed fresh,were dried and traded to the Nass RiverTsimshian every spring for eulachon grease(Turner 1974). Dawson (1880), likelyreferred to sgyuu, as follows: “A sea-weedresembling dulse, but which I have only seenin dried cakes, is found, especially in thesouthern islands, preserved by drying andboiled into a sort of tea or soup.”

The Haida word for foliose lichens isihk’inxaa-kwii7aawaay [“forest-cumulus-cloud”], but relatively few Haida nameshave been recorded for lichens and fungi(Turner 1974).

METHODS

“Many regions of conservation concern areincreasingly threatened by environmentaldegradation, and conservation biologistsoften require copious quantities of dataconcerning the distribution of species inorder to establish conservation priorities.”Snow and Keating (1999)

For the species lists we will eventually adoptthe nomenclature and systematics of theIntegrated Taxonomic Information System(ITIS). The ITIS is a dynamic biologicalchecklist supported by government agenciesof Canada, the U.S. and Mexico andmaintained by a network of over 1000taxonomists. It is accessible through theWorld Wide Web (in Canada: http://res.agr.ca/itis and in the U.S.: http://www.itis.usda.gov/plantproj/itis/index.html).

The ITIS conforms with the InternationalCodes of Botanical and ZoologicalNomenclature. Its purpose is to provideaccurate, scientifically credible and currenttaxonomic data standards for the comparisonof biodiversity datasets. Having stated that,ITIS can be expected to lag behind the mostrecent literature. The ITIS assigns a uniquecode (“Taxonomic Serial Number”) to eachtaxon and all attendant systematic levels upto Kingdom. The ITIS also includes, for allsystematic levels, taxon authorities,synonyms and data quality indicators.Central to the data quality process is peerreview prior to incorporation into ITIS, andperiodical review thereafter. The ITIS,therefore, facilitates international sharing,transfer and comparison of information.This is important for national agencies suchas Parks Canada when cooperating withinternational programs and global issues ofinformation sharing in support ofbiodiversity knowledge.

The ITIS is a work-in-progress and is not yetcomplete for some groups such as algae.Currently, the ITIS contains algae andlichens only from the U.S. Department ofCommerce, National Oceanic andAtmospheric Administration’s NationalOceanographic Data Center and these dataare not yet fully quality-controlled.Therefore, the algae in this report follow themost recent version of the University ofBritish Columbia (UBC) PhycologicalHerbarium master list as will be reflected inGabrielson et al. (in press). This list is theregional benchmark and is likely to beadopted by ITIS, as is Qian and Klinka(1998). For lichens, the nomenclaturefollows Qian and Klinka (1998) wheneverpossible. The ITIS is used for lichensystematics and for taxa not listed in Qianand Klinka (1998). Vascular plants(seagrasses) listed here are based on the

7

ITIS. When available, we will retrofit ourspecies lists with fully quality-controlledITIS codes.

Four major Pacific regional marine herbariaand the Canadian Museum of Nature (CMN)were contacted to establish the extent oftheir material from Haida Gwaii (AppendixA). The two institutions with material fromHaida Gwaii were the University of BritishColumbia (2758 records) and the CMN (9records).

Our intent was to capture all possiblesources and enter data into our GIS. Thestarting point was the Biophysical Inventoryof Coastal Resources (Harper et al. 1994)with its 107 shore observation stations inGwaii Haanas illustrated in Figure 2. Wethen looked back in time to marine

biological expeditions by universities andgovernment agencies for all of Haida Gwaii.Figures 3 to 5 contain the sample sites fromother noteworthy marine surveys. Figure 3contains sample sites in Gwaii Haanas fromtwo surveys commissioned by EnvironmentCanada-Parks before the establishment ofGwaii Haanas. The marine botanicalexpeditions to Haida Gwaii were theUniversity of British Columbia (UBC),Department of Botany synoptic surveys of1963 and 1976 supported by ProfessorRobert F. Scagel, with the latter publishedby Hawkes et al. (1978). Sample sites fromthese surveys comprise the majority of sitesin the UBC herbarium records illustrated inFigure 4. The sample locations from otherselected surveys of Haida Gwaii areprovided in Figure 5. Reports such asAdkins (1977) that have general regional

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Figure 2. The 107 shore observation locations from which marine plant data were recorded in the coastal biophysical inventory of Gwaii Haanas by Harper

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Figure 3. Sample locations from surveys of Gwaii Haanas commissioned by Environment Canada -Parks.

8

species lists, but with no specific localitydata, were not included.

Marine plant species data quality can beranked as follows:

1. species for which there are herbariumspecimens;

2. species mentioned in internationally peer-reviewed publications; and

3. species mentioned in “grey” literaturereports and unpublished surveys.

Within the grey literature, the expertise ofinvestigators determines data quality. Iftaxonomic specialists were engaged, thisincreases the report’s reliability.

RESULTS AND DISCUSSION

SPECIES LISTS

“The difficulty in defining biodiversity is inproving the absence of a species. Absenceof proof is not proof of absence.” Druehl(1999 a)

With very few exceptions, the historicalspecies listings were made without protocolsspecifically establishing the absence ofspecies. Lists of seaweeds and seagrassesand marine lichens in the Gwaii Haanasdatabase are provided in Appendix B.Included are the number of sites at which allmarine plant taxa have been reported fromHaida Gwaii (Queen Charlotte Islands) andGwaii Haanas National Park Reserve. Alsoincluded are the number of voucheredspecimens for each species at the UBC orCMN herbaria and the map number for thedistribution of that species illustrated inAppendix C. Part 1 of Appendix B coversmarine algae and seagrasses and Part 2covers marine lichens, all of which arevouchered CMN specimens.

Algae

Algal species names in Appendix B arebased on Keys to the benthic marine algaeand seagrasses of British Columbia,Southeast Alaska, Washington and Oregon[Revised Edition] (Gabrielson et al. in press)that reflect the most complete and currentupdate of the Scagel et al. (1993) Synopsis.The important exception is the exclusion of10 new species combinations that will beissued first in print by Gabrielson et al. (inpress). Their names (to be replaced) aretagged in Appendix B to alert readers to theimminent change.

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kept in the UBC Herbarium.

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Figure 4. Sample locations in Haida Gwaii from whichmarine plant specimens are retained at the University of British Columbia Phycological Herbarium.

9

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A Queen Charlotte Survey (TEC, 1992)B Smith and Buttler (1998)C Murchison-Faraday Passage Survey (TEC/HFP, 1994)D Burnaby Narrows Survey (TEC/HFP, 1993)E Rose Harbour Survey (AMR, 1997)

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Figure 5. Sample locations of various marine surveys in Haida Gwaii from which marine plants weredentified.

10

Concerning range extensions since Scagel etal. (1993), excluding name changes, theonly two extensions are for Cirrulicarpussp. nov. and Myriogramme repens, both ofwhich require checking with originalmaterial (M. Hawkes, personalcommunication). We underscore that thereremain significant seaweed taxonomic andsystematic problems whose resolutions willchange the region’s marine flora appreciablyin the future.

Table 2 lists the nine marine algal specimensat the Canadian Museum of Nature. Allspecies are listed in Appendix B, but onlythe two, those from Rennell Sound, aremapped in Appendix C, described below.

Haida Gwaii is of inherent marinebiogeographical interest as it represents thesouthern extreme of the northern seaweedflora (Druehl 1981, 1999 a). Haida Gwaiihad 32 species not reported from northernBritish Columbia mainland coasts (Hawkeset al. 1978). Garbary et al. (1980) addedanother 16 to this count, but also mentionedthat over 50 taxa known from Alaska andsouthern British Columbia were not

recorded from northern British Columbia.Garbary et al. (1980) commented that suchfindings could be due to insufficientcollecting from northern British Columbiacoastal areas. The four floristic categoriesof Hawkes et al. (1978): northern species(found north of 59o N), southern species(found south of Washington State), localspecies (occurring between 59o N andWashington State only) and cosmopolitanspecies (occurring north and south along theentire coast), occurred in equal proportionsfor the northern mainland and Haida Gwaiicoasts. Within Southeast Alaska (Icy Bay[~60o N] to Dixon Entrance) alone, O’Clairet al. (1996) identified three seaweed areas.They found that “outer coast” (warmer,higher salinity), “northern inside waters”(colder, lower salinity) and “southern insidewaters” (warmer, lower salinity) sharedmany species, but each also had distinctspecies. Perhaps Haida Gwaii comparesmost closely with the “outer coast” typearea. Clearly, much more collecting needsto be done in the northern British Columbia-Southeast Alaska region.

With focused marine botanical effort, theseaweed species diversity recorded withinGwaii Haanas will likely increase as currentknowledge represents ~49% of regionalseaweed biodiversity. Table 3 lists thenumber of taxa recorded from Gwaii Haanasand Haida Gwaii as a whole. The seaweedtaxa of Haida Gwaii in Appendix B excludethe “unidentified” specimens in thedatabase. In Table 3 we did include lonegenera for which no species were assigned.The Oregon to Southeast Alaska regionlistings are based on Scagel et al. (1993),Hawkes (1992, 1994 a), Tunnicliffe (1993)and Gabrielson et al. (in press).

* no Latitude or Longitude recorded

Table 2. Herbarium marine specimens from Haida Gwaii in the Canadian Museum of Nature, Ottawa.

Species (current name)[National Herbarium No.] Location* Collector (Year)

Spongomorpha coalita(Acrosiphonia coalita) [4618]

QCI Spreadborough (1911)

Ulva lactuca latissima(Ulva fenestrata) [4347]

QCI Spreadborough (1911)

Ralfsia verrucosa(Ralfsia pacifica) [4614]

QCI Dawson (1878)

Ceramium rubrum [4620] QCI Spreadborough (1911)Rhodomela lycopodioides [4485] QCI Spreadborough (1911)

Bossiella sp. [5189] Rennell Snd. W.v.V. (1966)Corallina sp. [5188] Rennell Snd. W.v.V. (1966)

Cryptosiphonia woodii [4617] QCI Spreadborough (1911)Rhodymenia pertusa

(Sparlingia pertusa) [4585]QCI Spreadborough (1911)

11

Seagrasses

The following three seagrass species areconfirmed to occur in Haida Gwaii waters:

• “eelgrass” (Zostera marina) found alongsheltered sediment shores; and

• two “surfgrasses” (Phyllospadix scouleriand P. torreyi) growing from bedrockcrevices and among boulders and cobbles ofexposed rocky shores.

There was a consultant’s report originatingfrom an assessment for the new harbour inSandspit recording a thin-bladed species asthe introduced species Zostera japonica,which is now common in the southern Straitof Georgia. Those specimens, plus somefrom Queen Charlotte City collected in 1999by us, were confirmed by Dr. Paul G.Harrison of UBC to be Z. marina (P.G.Harrison, personal communication).

There is an anomaly in the form of Ruppiamaritima which is illustrated in Pojar andMacKinnon (1994) as occurring throughoutHaida Gwaii. There are herbariumspecimens as documented in Calder andTaylor (1968), but there are no records fromthe marine herbaria we contacted. Pojar and

MacKinnon’s (1994) map is likely based onCalder and Taylor’s (1968, p. 70)generalizations on the occurrence of R.maritima (along with Z. marina further toseaward) in Haida Gwaii estuarine saltmarshes. The species is not discussedfurther here. Regional seagrass specialiststend not to view R. maritima as a seagrass,but as a brackish water grass (Wyllie-Echeverria and Thom 1994), although thespecies is also assigned to the upperintertidal (Gabrielson et al. 1990).

Although three species of seagrasses occurin Gwaii Haanas, they likely have adisproportionately important impact onbiodiversity in shallow marine areas becauseof the communities of organisms associatedwith them. The federal-provincial MarineProtected Areas strategy for the Pacificidentified the limited amount of eelgrasshabitat as areas of “primary concern” forprotection (Anonymous 1998, p. 9).

Marine Lichens

The 88 marine lichen species listed inAppendix B (Part 2) all represent voucheredspecimens held in the Canadian Museum ofNature. The substrate type and shorezone(s) occupied by all species are included.

* 34 species are known only from B.C. Waters

Table 3. Seaweed and seagrass taxa of Haida Gwaii and the Oregon to Southeast Alaska region.

Algae / SeagrassType

GwaiiHaanas

HaidaGwaii

Whole Region

GeneraSpecies/

Subspecies

Red (Rhodophyta) 151 225 161 373Brown (Phaeophyta) 46 73 66 143Green (Chlorophyta) 22 45 51 171

Golden (Chrysophyta) 1 1 3 6Seagrass (Anthophyta) 3 4 3 6

Totals 223 348 284 645*

Algae / SeagrassType

GwaiiHaanas

HaidaGwaii

Whole Region

GeneraSpecies/

Subspecies

Red (Rhodophyta) 151 225 161 373Brown (Phaeophyta) 46 73 66 143Green (Chlorophyta) 22 45 51 171

Golden (Chrysophyta) 1 1 3 6Seagrass (Anthophyta) 3 4 3 6

Totals 223 348 284 645*

12

The zonation designations are preliminary.

GEOGRAPHIC DISTRIBUTION

Appendix C comprises individual mapsillustrating the distribution of every speciesof alga and seagrass listed in Appendix B(Part 1). The species are illustrated in thesame order that they are listed in AppendixB. The map number from Appendix C iscross-referenced with each species inAppendix B (Part 1).

Algae

As a first approximation of kelp forest

distribution, Figure 6 shows the occurrenceof kelp symbols from CanadianHydrographic Service charts (3807, 3808,3809, 3825, 3827, 3894) of the MoresbyIsland area, including much of GwaiiHaanas. The kelp species illustrated aremost likely to be those forming conspicuousforest, such as giant kelp (M. integrifolia)and bull kelp (Nereocystis luetkeana), whichare visible as floating kelp forest canopyalong rocky shores. Note that the west coastof Gwaii Haanas north from Nagas Point toTasu Sound is uncharted; hence no kelpsymbols. Kelp is likely ubiquitous in GwaiiHaanas and could be expected to occur alongall rocky shores.

Illustrated in Figure 7 is the inferreddistribution of kelp in Gwaii Haanas fromthe Biophysical Inventory (Harper et al.1994) aerial videotaping. Kelp distributionwas derived largely from the videotapes;with some verification from the 107 shoreobservation stations illustrated in Figure 2.

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Legend

Canadian Hydrographic ServiceChart Boundaries with chartnumbers indicated.

#

Kelp locations as indicatedon Canadian HydrographicService charts.

##

#

3857

0 25 50 75 100 Km

132 W133 W 131 W

52 N

53 N 53 N

54 N 54 N

131 W

Figure 6. The distribution of kelp symbols (likely representing mostly and

) on six Canadian Hydrographic Service nautical charts of the Moresby Island area.

Macrocystis integrifolia Nereocystis luetkeana

LegendKelp Forest

0 25 50 Km

132W 131W

52 N52N

132W 131 W

Figure 7. Inferred distribution of kelp (likely dominated by and

) from the coastal biophysical inventory of Harper (1994).

Macrocystis integrifolia Nereocystis luetkeana

et al.

13

The largest kelp forest in southern HaidaGwaii is on Fairburn Shoals, CumshewaInlet mapped by Haegele and Hamey (1981)for herring spawning area surveys. Otherextensive kelp forests are located along thenorth shore of Graham Island (Coon et al.1979).

Seaweeds and seagrasses are usefulindicators for assessing wave exposurecriteria of rocky shores. Table 4 listsoccurrence of selected plant species used todiscriminate community types according toexposure, substrate and tidal zone in GwaiiHaanas by Harper et al.(1994). This wasrefined by Druehl and Hopkins (1999) intheir survey of SGaang Gwaii shown inTable 5 and illustrated in Figure 8, in whichboth have the Harper et al. (1994) exposureclassifications. The presence of Mazzaellacornucopiae and Zostera marina in Table 5bracket the full exposure range within whichthere are at least four core species perexposure class. Some of the exposureclassification differences in Figure 8, such assites P and R, are large and likelyattributable to transcription errors in the1994 biophysical inventory. This is notsurprising, given the large scale of thatproject. Other differences are refinementsdue to better information on the presence ofindicator species. Druehl and Hopkins(1999) speculated that, unlike the mainland,rapid seawater mixing and less freshwaterrunoff from the limited land mass means thatsubstrate, wave exposure and temperaturedetermine kelp distribution with littleimpacts from salinity. This could extend therange of wave exposures at which somespecies may be found in Haida Gwaii asspecies do not experience as much salinityfluctuation.

In 1999, a preliminary seaweed survey wasdone in the Ellen Island region. The six

permanent intertidal seaweed monitoringtransect sites and the reference and aberrantMacrocystis sp. sites are shown in Figure 9.There is no transect site No. 3. The transectswere established based, in part, on a protocoldeveloped for Pacific Rim National ParkReserve by Druehl and Elliott (1996). Theobjective of the protocol was to identifycommunities based on kelp assemblage-shore physical attribute correlations.Monitoring the rich kelp assemblages fromdifferent shore exposure types of GwaiiHaanas over time has benefits as discussedbelow.

Besides the aberrant giant kelp being anoteworthy botanical anomaly, its presenceis an issue of potential interest to the herringspawn-on-kelp fishery as discussed below.The northern giant kelp (M. integrifolia)occurs from Sitka, AK to Monterey, CA(Foster and Schiel 1985). Until recently, thesouthern giant kelp (M. pyrifera) wasreported from Monterey, CA to BajaCalifornia, Mexico (Foster and Schiel 1985),but O’Clair and Lindstrom (2000) havereported it from Alaska waters based on anold Setchell and Gardiner report. Thespecies’ preference for moderate waveexposure is well known (Druehl 1978). InGwaii Haanas, giant kelp normally occurs onmore sheltered shores than bull kelp (N.luetkeana) and is found shoreward of bullkelp where the two coexist (e.g., at the“reference” sites in Figure 9). South ofBenjamin Point on Moresby Island,however, giant kelp was found seaward ofbull kelp along exposed shores (e.g.,“aberrant” sites in Figure 9). The identity ofthis giant kelp type cannot be resolved untilholdfasts, diagnostic features of Macrocystisspp., can be retrieved for examination. Thesouthern giant kelp is larger (to ~45 m) andgrows at deeper sites than the northern giantkelp (to ~10 m).

14

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Legend

* Wave exposure classification from Harper (1994): VE = very exposed; E = exposed; SE = semi-exposed; SP = semi-protected; P = protected.

et al.

** Revised “Harper” exposure classification by Druehl and Hopkins (1999)

- = no change from the original “Harper” exposure classification.

S aangGwaiiG

Nan SdinsVillage Site

131°13’ W

52°5’ N

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52°6’ N

0 500 1000 1500 Metres

#B

Figure 8. Locations of the 19 wave exposure assessment sites in the S aang Gwaii archipelago surveyed in July, 1999 by Druehl and Hopkins (1999).

G

16

The only invasive seaweed recorded so farin Haida Gwaii is Sargassum muticum. It isspeculated to have been introduced into theStrait of Georgia and Puget Sound, WA inthe 1930s from imported Japanese oysters.The species now ranges in the NortheastPacific from Baja California, Mexico northto southern Southeast Alaska (O’Clair et al.1996). Figure 10 illustrates the distributionof S. muticum in Haida Gwaii. Included areobservations made on the south side of EllenIsland in July, 1999 (Druehl and Hopkins1999) and just south of Lawn Hill (Ms. M.

Stronge, Gwaii Haanas, personalcommunication).

Seagrasses

Figure 11 illustrates the distribution ofeelgrass (Z. marina) which prefers sheltered,sedimentary shores and surfgrasses(Phyllospadix spp.) which tend to grow onexposed, rocky shores. Overall, there is atendency for surfgrasses to be recorded frommore exposed sites. There are anomaliesthat require verification such as surfgrass(not eelgrass) reported from Masset Inlet.

##

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52°10' N52°10' N

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MacrocystisMacrocystis

Area ofInterest

Figure 9. Locations of the six permanent intertidal seaweed monitoring transects and sp.observations made in July, 1999 by Druehl and Hopkins (1999).

Macrocystis

Table 5. The expected presence/absence of marine plant species indicating wave exposure on S aang Gwaii.GExposure

Class*Mazzaella

cornucopiaeAlarianana

Lessoniopsislittoralis

Laminariasetchellii

Nereocystisluetkeana

Laminariabongardiana

Macrocystisintegrifolia

Fucusgardneri

Zosteramarina

VE + + + + + - - - -E - + + + + - - +/- -

SE - - + + + +/- - + -SP - - - +/- +/- +/- + + -P - - - - - +/- +/- + +

* Exposure classes according to Harper (1994): VE = very exposed / E = exposed / SE = semi-exposed / SP = semi-protected / P = protected+ = expected present / = expected absent

et al.

-

17

Within Gwaii Haanas, 254 sites fromvarious surveys were ordered according totheir shore exposure category as defined in

the biophysical inventory (Harper et al.1994). Listed in Table 6 is an examinationof these exposure categories of the sites atwhich eelgrass and surfgrass occurred. Thesample size for the “very exposed” sites issmall, but the “exposed” and “semi-exposed” sites show particularly well theconcurrence of the shore exposure ratingwith the expected differing exposurepreferences of each species. This revealsthat the current state of the Gwaii Haanasdatabase, with its commitment to the shoreexposure classification system, reasonablyreflects habitat preferences of some species.

In 1999, Gwaii Haanas’ Warden Servicesbegan an intermittent task of mapping theeelgrass meadows which straddle the lowerintertidal to the shallow subtidal. Illustratedin Figure 12 is the first output from thislong-term project. It is likely that Z. marinais ubiquitous along sheltered sedimentaryshores of Gwaii Haanas. On-site mappingcould also provide ground-truthing forpossible future remote sensing of GwaiiHaanas’ nearshore resources.

Distribution maps are viewed as importantfor establishing seagrass baselines (Wyllie-Echeverria and Thom 1994). This is doublyimportant for Gwaii Haanas as eelgrass areassociated with sheltered sedimentary shores

#

#

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##

54 N

132 W

54 N

131W

132 W133W

53 N53 N

52 N

Legend# Sargassum muticum

0 25 50 75 100Km

Figure 10. Distribution of from various marine surveys in Haida Gwaii.

Sargassum muticum

Shore Exposure CategoriesVery

Exposed ExposedSemi-

ExposedSemi-

Protected ProtectedTotal Sites 20 54 73 61 65

Zostera marina 2 (10%) 0 (0%) 6 (8%) 10 (16%) 26 (40%)Phyllospadix spp. 6 (30%) 25 (46%) 31 (42%) 19 (31%) 8 (12%)

Total Sites

Notes:The number of sites by exposure category according to Harper (1994) is based on a spatial proximity analysis that tallies samples if they are within 100 m of a given coastline class. Using this methodology, it is possible for a sample to be tallied more than once because it can be within 100 m of more than one coastline class. For this reason, the 273 total for the top row is greater than the actual number of 254.

et al.

Table 6. The number (and percent) of sites at which seagrasses occurred according to exposure category at 254 sites in Gwaii Haanas.

18

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Phyllospadix#

Zostera marinaS

0 50 100 Km

Legend

spp.

Figure 11. Distribution of eelgrass ( ) and surfgrasses ( spp ) from various surveys in Haida Gwaii.

Zostera marina Phyllospadix .

19

which, in the case of estuaries, representonly ~6% of Gwaii Haanas’ shoreline(Harper et al. 1994). Yet, estuaries areundoubtedly important to the juvenilesalmonid production of Gwaii Haanas aswell as being nurseries for other inshore fishand invertebrates.

MARINE PLANT ISSUES IN GWAIIHAANAS MANAGEMENT

“…. the most critical part of maintainingbiodiversity is maintaining the dynamic setof interactions that define what thatecosystem is and how it functions.”(Suchanek 1994)

“The protection of natural, self-regulatingmarine ecosystems is important for the

maintenance of biological diversity”Preamble: Marine Conservation Areas Act

Marine plants and their associatedcommunities are immensely important to thenearshore ecosystem structure and functionin Gwaii Haanas. Inventory, in isolation, isof limited interest to managers unless it isplaced in a broader context. Inventory is theacknowledged first step in takingresponsibility for an area’s marinebiodiversity (Hawkes 1992; Suchanek1994). Maintaining biodiversity is acornerstone of ecosystem structure andfunction - the general goal of ecosystem-based management (Grumbine 1994). Themost stringent conservation norm has beensuggested by Callicott et al. (1999) as beingbiological integrity, defined as; “… nativespecies populations in their historic varietyand numbers naturally interacting innaturally structured biotic communities.”This includes retaining biological elements(i.e., biodiversity) and processes. Marineinventories are, therefore, linked toecosystem-based stewardship as articulatedin the proposed Marine Conservation AreasAct.

Kelp forests and seagrass meadows providethe following ecosystem functions:

• inputs detritus into coastal food webs(Phillips 1984; Duggins et al. 1989);

• yields a “trophic subsidy” of plantbiomass from the subtidal to intertidalgrazers (Bustamante et al. 1995);

• provides energy from beached drift algaland seagrass wrack (Polis and Hurd 1996);

• moderates nearshore water movement(Phillips 1984; Lobban and Harrison 1997);

Figure 12. An example of eelgrass ( )mapping in Louscoone Inlet, summer, 1999.

Zostera marina

LouscooneInlet

Legend

meadow

0 1 2 3Km

131°10' W

52°10' N

131°10' W

52°10' N

Areaof

Detail

Zostera marina

20

• forms a biogenic nearshore habitatarchitecture of nurseries of juvenile fish andinvertebrates (Phillips 1984; Foster andSchiel 1985);

• creates biogenic habitat for adults of manyspecies, including those of commercial andcultural value such as northern abalone(Haliotis kamtschatkana) (Sloan and Breen1989) and rockfish (Sebastes spp.)(Carr1994); and

• underpins the basis for traditional andcommercial Pacific herring spawn-on-kelpfisheries.

Concerning marine stewardship, theimportance of marine plants is manifested inthe following nearshore Gwaii Haanasmarine ecosystem issues that include:

• understanding the role of plants inecosystem structure and function, such as therole of seagrass meadows in the land-sealinkages in our estuaries;

• monitoring environmental threats to, andwell-being of, ecosystems;

• assessing impacts of repatriated sea otters(Enhydra lutris) on nearshore kelp forestecosystems;

• evaluating the role of red sea urchin(Strongylocentrotus franciscanus) grazing(herbivory) on kelp forest ecosystems;

• cooperating in the Department of Fisheriesand Oceans’ northern abalone stockrebuilding;

• maintaining healthy kelp resources in aidof Pacific herring SOK harvest; and

• assisting with backcountry monitoring of

visitor impacts to intertidal habitats.

Nearshore Marine EnvironmentalMonitoring

“Without reliable data about rates of changewithin any habitat, there is no possibilitythat we can predict the sorts of changes thatare likely to be associated with humaninterferences and developments.”(Underwood and Kennelly 1990)

“Long-term ecological monitoring is the firststep in learning how to assess ecosystemhealth.” (Davis 1993)

Monitoring is becoming a fundamentalcomponent of Parks Canada’s mandate ofsustainable management and use withoutcompromising the structure and function ofecosystems (Woodley 1993). Davis (1993)summarized the utility of the U.S. NationalPark Service’s marine monitoringcommitment as follows:

• indicating ecosystem health;

• defining limits of normal variation;

• identifying abnormal environmentalconditions; and

• verifying agents of abnormal change.

The process of monitoring developmentshould be experimental, begin with aconceptual model of the ecosystem and befocused on the population dynamics ofselected species relative to physio-chemicalenvironmental variables (Davis 1993).

At a workshop assessing ecological statushosted by Pacific Rim National ParkReserve, both kelp and seagrass wereselected as potential marine environmental

21

indicators of merit on the British Columbiacoast (Rowe et al. 1999). Marine plants areeffective sentinels for monitoring the well-being of inshore marine ecosystems andreflecting ocean climate changes.Monitoring can be done using both intertidalspecies and shallow subtidal species withinthe photic zone (~35 m depth limit of lightpenetration). Seaweed in the upperintertidal, kelp and seagrass in the lowerintertidal and shallow subtidal, and watersurface kelp forest canopy in the subtidal areall possible targets. Monitoring suites ofplant species with known environmentaltolerances can permit establishing potentialconsequences of environmental change(Druehl 1999 b). Both kelp forests andseagrass meadows in intertidal and shallowsubtidal waters are amenable to surveillanceby remote sensing (Deysher 1993;Guillaumont et al. 1993; Israel and Fyfe1996).

Without seasonal and interannual data ondistribution and abundance of algae, nocomparison between locations is possible(Underwood and Kennelly 1990). Theseauthors proposed the following two stagestowards understanding intertidal and subtidalrocky shore plant communities:

• quantitative studies to describe the patternsof distribution and abundance of algae withappropriate spatial and temporal replicationto estimate variances among and withinshores and among years and seasons; and

• experimental analyses of the processescausing the observed patterns.

The latter is the evolutionary end-point ofsuch work and has not yet been achievedanywhere in temperate coastal systemsworldwide.

The positive attributes of seaweed formonitoring are defined by Druehl (1999 b)from the Pacific Rim National Park Reserveworkshop as follows:

• stationary plants must integrateenvironmental effects or perish;

• have variable age structures (ephemeral toperennial) spanning duration of changes;

• occupy diverse nearshore habitats(intertidal/subtidal) and wave exposureregimes;

• possess variable physiological tolerances;

• convenient – abundant, large size, speciesrich, easy to locate and identify; and

• high degree of morphological plasticityreflecting environment conditions such aswave exposure.

Intertidal Seaweed and Subtidal KelpForest

“…. an extensive monitoring program forinshore macroalgal habitats is a valuabletool for evaluating effects of environmentalchange.” (Milligan et al. 1999)

“ …. analyzing community patterns formarine algae via groupings based onfunctional aspects of their morphology andanatomy provides substantial insight intocommunity structure.” (Steneck and Dethier1994)

“ …. no matter how well one understandskelp populations, any current program willfail to discern the ghosts of missinganimals.” (Dayton et al. 1998)

The National Park Service in California has

22

been the leader in long-term intertidalmonitoring in Northeast Pacific NationalParks. Monitoring for general intertidalecosystem status began in Channel IslandsNational Park in 1982 (Richards and Davis1988) and the first long-term visitor impactassessment was at Cabrillo NationalMonument in 1990 (Engle and Davis 1996).

A summary of plant components of someintertidal marine monitoring initiatives fromAlaska to California is provided in Table 7.Plants clearly play a prominent role inintertidal monitoring. Note how recent theseinitiatives are. This is a time of growth andwidespread acceptance of the need for long-term intertidal monitoring in National Parks.

Intertidal monitoring is usually focused ondiscrete bands characterized by dominantspecies, among which bands of vegetationare prominent. For example, Richards andDavis (1988) monitored their upper to midintertidal rocky shores including vegetatedbands described in Table 7. In BritishColumbia, “bio-band” codes based, in part,on conspicuous bands of certain plant typesare used in zonation mapping schemes formarine shores (Searing and Frith 1995) andestuaries (Howes et al. 1999). A detailedreview of rocky intertidal marine monitoringis in draft form for the California Sea GrantProgram (Murray et al. in press).

In Barkley Sound, off the west coast ofVancouver Island, Milligan et al. (1999)used intertidal kelp populations ofHedophyllum sessile to document effects ofthe 1997-1998 El Niño. They drew upon thearea’s only long-term (1991-1998) kelp datato evaluate the negative (poor growth due todepressed nutrient levels) impacts of thatwarm-water event. Among the variablesmeasured were percent cover, adult-juveniledensity, plant reproductive status, biomass

and cover of understory species.

Of all rocky nearshore subtidal habitats inthe Northeast Pacific, kelp forests dominateecosystem architecture and are the mostinvestigated. There are sufficient time seriesdata from which some ecosystemgeneralizations can be made. Kelp forestaerial surveillance began off the southernCalifornia coast in the 1930s with regularsurveys by the commercial kelp harvestsector since the 1950s (Deysher 1993).Regular aerial scientific broad-scale surveysbegan in 1967 (North et al. 1993) andquarterly in situ surveys by divers at PointLoma, near San Diego began in 1983(Tegner et al. 1997). Kelp forest monitoringhas also been undertaken at Channel IslandsNational Park annually since 1981 (Davis1988; Davis et al. 1994; Davis et al. 1996;Davis et al. 1997). Also in the area, SPOTHigh Resolution Visible (HRV) digitalmultispectral (XS) satellite images wereevaluated for their use in mapping andquantifying giant kelp forest (Augenstein etal. 1991). The authors concluded that SPOTHRV-XS digital data yielded kelp surveymaps sufficient for resource mapping andmonitoring and that imagery closelycorresponded to distributions derived fromcolour infrared aerial photographs. No otherregion of the Northeast Pacific has acomparable long-term kelp monitoringhistory as southern California.

Detecting kelp ecosystem trends dependsupon two things (Dayton et al. 1998):

• a good description of the benchmark (i.e.,a sound baseline/inventory); and

• an ability to establish distinctions betweennatural and human changes.

There is doubt, however, as to what kelp

23

Table7.Sum

mary

ofselectedintertidalm

onitoringattributes

involvingm

arineplantsin

NationalParksand

Monum

entsfrom

Alaska

toCalifornia.

Park(Proponent)

StartDate

Frequency*(m

o/season)Notesontheprotocolsand

plantspeciesReference

GlacierBayNationalPark

andPreserve(N

PS/USG

S1)

1997Annual(June)

Uses200mlongshore

segmentsforcoarse-grainedtransect/quadratsurveysanda

subsetofthese

forfine-grainedtransect/quadratsurveys;lichenVerrucaria

spp.usedto

demarcatelandward

sideofsegm

entsandFucusgardneritheonly

plantsam

pled(%cover,frond

lengthand

re productivestatus)butm

entionedsevenothergeneraas potentialm

onitoringcandidates

Irvine(1998)

PacificRim

NationalPark

Reserve(ParksCanada)1997

Annual(June)

Modified

theprotocolsoftheD

FO2Shorekeepers’M

anual;1m2m

idintertidalquadratsareusedin

differentrockyhabitats(tide pool/bench/surgechannel/vertical

face)examined

forallconspicuousplantandanim

alspecies,numberand

%cover

byvolunteer participants

Holmes(1999)

Olym

picNationalPark

(U.S.NationalParkService)1988

Biannually3

(summ

er)M

ethodrevised

in1997aftersampling

experimentsin1996;

quadrats(20X20cm

)sam

pledalong

20mtransectswithin

3recom

mendedshore

levelsdownto

themid

rockyintertidal;plantsre present~50%of“good

“taxaformonitoring

atalllevels

Dethier(1997)

ChannelIslandsNationalPark(U.S.NationalParkService)

19824

Annual(January)

Permanentphotoquadrats(50X75

cm)recording%

cover:withinanupperm

ostbandofred

turfalgae( Endocladiamuricata),abandoffucoid

rockweed

(Pelvetiafastidata )andHesperophycusharveyanus,thena

bandofredalgae

(Gigartinasp.and Gelidium

sp.)justlandward

ofmussels

Richards&Davis(1988)

Davisetal.(1994)

CabrilloN

ationalMonument

(U.S.NationalParkService)1990

Semi-annual 5

(spring/fall)Perm

anentphotoquadrats(50X75cm)recording

%cover;fucoidrockw

eed(Pelvetia

fastidata)/10m

transectcounts;Egregiamenziesii,Sargassum

muticum,Phyllospadixspp.,Corallina

spp.

Engle&D

avis(1996)

*actualorrecomm

ended1

TwoU

.S.DepartmentofInterioragencies(NationalParkServiceand

U.S.GeologicalSurvey’sA

laskaBiologicalScienceCenter),developedthismethod

foratotalof

threeparks,includingalso:K

atmaiNationalPark

andPreserve

andW

rangell-St.EliasNationalParkandPreserve

2Departm

entofFisheriesandOceansdeveloped

its Shorekeepers’Manual(Jam

iesonetal.1999)3

After1993therew

aseven-yearandodd-yearsam

plingdividedam

ongthe22

rockyshorelocations

4AnacapaIs.in

1982,expandedto

15locationsamongthe

islandsby19885

Threesitesatonelocation

24

forest monitoring can achieve given the poorhistorical baseline from undisturbed kelpforests and the inadequate understanding ofhuman (e.g., harvest and pollution) impactson kelp forest ecosystems (Dayton et al.1998). From 25 years of kelp forestobservations in southern California, Daytonet al. (1998) concluded that the definition ofa meaningful benchmark was “impossible”because:

• many large kelp forest predators have beenlocally extirpated for decades;

• kelp are very susceptible to irregular,large-scale, low-frequency oceanographicphenomena such as warm water, low-nutrient El Niño events (Tegner et al. 1997)and episodic severe storms (North et al.1993); and

• fisheries still have major, but poorlyunderstood, impacts on kelp forestecosystems.

In the wilderness area comprising GwaiiHaanas, fishing impacts on kelp forests arelikely to be much less than those in southernCalifornia. With the significant exception ofsea otters, the suite of large kelp-associatedpredators in Gwaii Haanas is relativelyintact. Nonetheless, we cannot differentiatebetween natural and human impacts on localkelp forest ecosystems. We know nothingabout how ocean climate affects local kelpforests, nor do we know about indirectcommercial harvesting impacts.

On the one hand, kelp are useful formonitoring physiochemical attributes ofocean climate such as temperature andnutrients (Tegner et al. 1997; Druehl 1999a,b). Moreover, examining algae yieldsinsight into environmental conditionsbecause, when observed at the functional

group level, they are temporally stable andpredictable (Steneck and Dethier 1994).This is especially applicable compared tostudying algae at the species level. On theother hand, the generalization by Ward et al.(1999) that marine plant assemblages arepoor surrogates for overall species richness(i.e., biodiversity) may apply to kelp forestsas an assemblage - but perhaps not as ahabitat. Firstly, because of the patchiness ofkelp forests that includes species richness ofnon-vegetated habitats. Secondly, becauseof the suggestion by Dayton et al. (1998)that kelp forest communities can be looselyintegrated with weak mutual dependencies,e.g., large predatory fish can be removedwith little effect to the rest of the ecosystem.Perhaps the solution is functional group(total kelp forest) focus on both canopy andunderstory floras and attendant faunas aschanging environmental conditions favordifferent kelp forest components.

In summary, we do not yet know enoughabout overall kelp forest status to reflectnearshore ecosystem structure and functionin Gwaii Haanas. Kelp forests haveattributes worth investigating, but a long-term commitment would be needed tounderstand natural impacts such as thoseinduced by ocean climate (temperature,nutrients and waves) and a better knowledgeof human impacts dominated by commercialfishing.

Reports are just now emerging on responsesof seaweed populations in “no-take”reserves. Edgar and Barrett (1999)recounted from temperate Tasmanian reefsthat seaweed species richness increased andindividual species dominance changed inprotected versus reference areas.

25

Seagrass Meadows

Seagrass monitoring straddles the midintertidal through to the shallow subtidal.Seagrass meadows are amenable to remotesurveillance (aerial and satellite) and toground-truthing by various forms of in situdiver-related techniques (Kirkman 1990). Aconservation strategy suggested for seagrassmeadows stressed protection of discretemeadows within a larger area (McNeill andFairweather 1993). The greater range ofhabitat types, locations and aspectsassociated with different small meadowssheltered more biodiversity than a singlelarge meadow. As with kelp forests,connections among habitats and knowledgeof recruitment processes should be wellunderstood before protected site selectionand boundary delimitation.

In British Columbia, there is no regularseagrass monitoring (Colin Levings,personal communication). There is oneresearch series of aerial infra red and colourphotography of seagrass meadows nearbyRoberts Bank port on the Fraser River deltasystem (Tarbotton and Harrison 1996).Elsewhere in the northwest U.S. there are nolong-term seagrass monitoring sites (RonThom, personal communication). Remotesensing of seagrass meadows in Puget Soundwas abandoned by the Washington StateDepartment of Natural Resources becausepoor water clarity obscured their deeper,seaward portions (Ron Thom, personalcommunication). Washington and Oregonare collaborating in the Pacific NorthwestCoastal Ecosystem Research Study(PNCERS) to monitor, among other things,seagrass meadows for up to four years inWillapa Bay (WA) and Coos Bay (OR)starting in 1998.

Remote sensing using, for example,

multispectral SPOT XS imagery to monitorwell-being and distribution of seagrasses iswell known (Kirkman 1990; Israel and Fyfe1996). A variety of intertidal and shallowsubtidal cover types can be differentiated.Remote sensing coupled with GIS isparticularly efficient for data collection andanalysis, and provides good potential formonitoring applications. The water clarityaround Gwaii Haanas is amenable to remotesensing but the weather is often overcast.The database being developed by theWarden Services could provide a baselinefor comparison with remote imagery.

Global threats to seagrasses come fromhuman modification of watershedsdecreasing downstream coastal waterquality, other coastal pollution and coastaldevelopment (Fong et al. 1997). For GwaiiHaanas, the undisturbed watersheds andundeveloped wilderness shoreline mean thatseagrasses are little threatened by humanactivities. Local seagrass meadowsrepresent, therefore, relatively undisturbedecosystems.

Marine Lichens

Outside of California, no marine lichenspecies has been used for environmentalmonitoring in the Northeast Pacific. Lichensare, however, well established as airpollution monitors.

Coastal marine lichens have been used asenvironmental air pollution monitors ofmarine aerosols. In southern California,Ramalina menziesii, which also occurs oncoastal trees in Haida Gwaii (I. Brodo,personal communication), accumulatesmetals (Boonpragob and Nash 1990).Marine lichens (Ramalina spp.) have alsobeen used in monitoring marine aerosolpollution in Europe (Roux et Sigoillot 1987).

26

In the context of Gwaii Haanas’ wildernesssetting, lichens could be sentinels of distant,Pacific Rim atmospheric pollution. Once themarine lichen flora of Gwaii Haanas is betterknown, perhaps candidate species could beselected for monitoring regional pollution ofmarine aerosols.

Kelp Forest Ecosystem Issues

Sea Otters and Haida Gwaii History

The extirpated sea otter casts a broad shadowover Haida Gwaii – biologically andculturally. In 1778, Captain Cook took seaotter pelts from Nootka Sound and the nextyear traded them in Canton at ~1800% profit(Gibson 1992). This precipitated a scramblefor sea otter pelts that was strongly felt in theearly post-contact times of the 1790s inHaida Gwaii. For example, British tradersacquired thousands of sea otter pelts for theChina trade from Chief Cuneah of Kiustavillage on the northwest tip of GrahamIsland (Robinson 1996). Desire for sea otterfur intensified early European contact,brought great material wealth (e.g., irontools) and social change to the Haida.Contact from trade eventually broughtdevastating diseases as well, such as thesmallpox that ravaged the Haida in the1860s. The sea otter trade rapidly lost vigorand by 1825 potatoes became the majorcommodity for the depressed Haida tradingeconomy (Acheson 1995). Despite thedecline in the sea otter trade, Europeantrading presence for other commoditiesremained entrenched coast-wide.

The sea otter and its prey the red sea urchinare on the family crest of Gwaii Haanas.The relationships between these species andtheir kelp forest habitat are vital socio-cultural and scientific issues for GwaiiHaanas. Human reintroduction of sea otters

to establish nearshore ecosystems reflectingpre-contact status and subsequent impacts toresident red sea urchin and northern abalonepopulations will be important among themany long-term stewardship issues to beresolved for Gwaii Haanas.

Kelp Deforestation, Sea Otter and Red SeaUrchin

“Local extinction of sea otters hascascading effects of broad influence in kelpforest ecosystems.” (Estes et al. 1989)

“The concept’s (of keystone species)potential significance to conservationbiologists is that it designates species thatexert influences on the associatedassemblage, often including numerousindirect effects, out of proportion to thekeystone’s abundance or biomass.” (Paine1995)

Among kelp forests along rocky, nearshoreBritish Columbia and Alaskan coasts, seaotters have a keystone community role(Breen et al. 1982; Estes et al. 1989; Watson1993; Estes and Duggins 1995). There issome disagreement, however, on theirkeystone status. In California, Foster (1990)suggested that sea otter importance wasoverrated because, outside of sea otter range,red sea urchins and kelp abundances werehighly variable with ~20% of rocky coastdeforested. Estes and Duggins (1995) agreewith this area estimation, but counter thatsuch deforested habitat patchiness is typicalthroughout the Northeast Pacific where seaotters are a regional “keystone” predator.

The extirpation of sea otters from HaidaGwaii at some time before 1929 (Watson etal. 1997) likely led to an explosion of theirherbivorous red sea urchin prey andattendant heavy grazing on kelp often

27

causing kelp deforestation and the formationof sea urchin-dominated “barrens”. Hawkes(1994 b) speculated that lower seaweedbiomass and diversity has occurred coast-wide in British Columbia due to sea otterextirpation from their historical range. Thefactors that determine red sea urchinabundance in Haida Gwaii are complex. Forexample, in the absence of sea otters off thenorthern Graham Island coast, densities ofred sea urchins and kelp vary widely and redsea urchins do not dominate the presence ofkelp at all localities (Jamieson and Campbell1995). Jamieson and Campbell suggestedthat depth and shelter from wave action arealso important factors in these species’distributions.

Sea urchin barrens occur as a light-colouredband (due to encrusting coralline algae onrocks denuded of fleshy algae) seaward ofthe linear fringe of kelp forests along muchof Gwaii Haanas’ rocky shores. Whereas~20% of Northeast Pacific kelp forest areasnot occupied by sea otters are denuded ofkelp, the amount of such habitat isappreciably greater in Gwaii Haanas.Approximately 50% of rocky coastline ofGwaii Haanas is characterized by thepresence of kelp forest (Figures 6 and 7),half of that also has urchin barrensimmediately to seaward (Harper et al. 1994).Therefore, the return of sea otters could beexpected to have a significant effect on theexpansion of kelp forest as the red sea urchingrazers are consumed by sea otters. Thiswould be rapid as sea otters expand at a rateof ~20% annually when reoccupying newareas within their historical range (Watson etal. 1997; Woodby et al. 2000). Theecological effects of greatly expanded kelpforests in Haida Gwaii would mean muchmore structured nearshore adult and nurseryhabitat with attendant population increases,more detritus into nearshore food webs and

more drift algal wrack on the beaches.

Another potentially important species to kelpforest ecosystems in the North Pacificworthy of mention is Steller’s sea cow(Hydrodamalis gigas). Approximately20,000 years ago, this large (~10 m bodylength) highly specialized, non-diving kelpgrazer lived in herds throughout the NorthPacific (Domning 1972). The species wasdiscovered in its last refuge, the Commander(Komandorskiye) Islands in the NorthwestPacific, and was extinct by 1768. Thespeculation is that maritime aboriginalhunters extirpated Steller’s sea cow frommost of its range by the time it wasscientifically described from Bering’sRussian expedition in 1741 (Estes et al.1989).

There is much speculation concerning theecological relationships between kelpforests, sea otters and Steller’s sea cow.Estes and Steinberg (1988) suggested thatkelp evolved in a North Pacific environmentin which kelp herbivory was at low intensitydue to sea otter predation on sea urchins.Domning (1989) disagreed, suggesting thatsea cow herds likely placed intense grazingpressure on kelp. Pitcher (1998) speculatedon a potential keystone role of Steller’s seacow in nearshore North Pacific habitatswithin the last few millennia. Anderson(1995) theorized that sea cows may havebeen “precariously balanced on sea ottercarnivory” by evolving as specializedsurface-feeders of preferred kelp speciesmaintained by sea otter predation on seaurchins.

One could speculate that kelp forests ofHaida Gwaii may have developed throughthe influences of two important mammalspecies, sea otters and Steller’s sea cow. Thereintroduction of sea otters may not,

28

therefore, yield a complete kelp forestecosystem for the archipelago if sea cowswere extirpated some millennia previously.

Estes et al. (1998) have shown that addingan apex predator (the killer whale Orcinusorca) to a kelp forest ecosystem under top-down control from sea otters in the centralAleutian archipelago had predictable effectson plant populations of intense herbivory byred sea urchins released from predation.Effects of commercial fishing and climatechange likely decreased offshore forage fishresources that, in turn, decreased offshorepinniped prey populations of killer whales.This caused switching by killer whales topredate on inshore sea otters and attendantinshore ecosystem changes in keeping withthe sea otters’ keystone role. This is areminder of ecosystem linkages by whichoffshore events can induce significantinshore changes.

Kelp Forest-associated Fisheries

“Unfortunately, although populations ofnumerous potentially important species havebeen recently reduced in kelp forestcommunities, usually little or nothing isknown of the ecological consequences ofthese reductions.” (Estes et al. 1989)

Important commercial kelp-associatedspecies or groups in Gwaii Haanas includesea urchins, abalone and rockfish. The issueof indirect fisheries impacts on kelp forestecosystems continues to vex managers.Knowledge of changing species interactionsand succession patterns under variousharvest regimes in kelp forests is poor(Dewees and Davies 1992). Relevant toGwaii Haanas, for example, is how does redsea urchin harvest effect kelp and northernabalone recruitment?

The close association of kelp forests to redsea urchins and northern abalone introducesanother significant future managementquestion - harvest refugia. Both red seaurchins and northern abalone are broadcastspawners for whom fertilization efficiencydeclines with adult densities. Distributioninto clumps of appropriate size and density iscritical to the design of the protection regimefor sea urchins and abalone (Quinn et al.1993; Shepherd and Brown 1993). The ideais that abalone stocks, for example, arecomposed of local populations linked bylarval dispersal into metapopulations(Shepherd and Brown 1993). Sizes ofrefugia depend on the dispersal ability oftarget species. Which areas are “sources”[contribute disproportionately large numbersof recruits] and which are “sinks” [receiverecruits but contribute little] of recruitsbecomes critical marine conservationmanagement information (Roberts 1998).Knowledge of the association of thesespecies with Gwaii Haanas’ kelp forests willbe an important long-term managementissue.

Kelp forest-associated species also compriseimportant political and cultural values. Forexample, northern abalone (this region’sonly abalone species) has been closed to allforms of harvest since 1990 with no relief insight due to chronically low stocks(Campbell 2000). This has had attendantnegative impacts to Haida culture(Richardson and Green 1989) and has, alongwith the SOK industry, created politicalmomentum to address kelp-associatedresources. Further, in April, 1999 thenorthern abalone became the first Canadianmarine invertebrate to be designated byCOSEWIC (Committee on the Status ofEndangered Wildlife in Canada) as“threatened” [i.e., “likely to becomeendangered if limiting factors are not

29

reversed”]. When Environment Canada’sproposed endangered species legislationbecomes law, this could intensify emphasison abalone conservation Province-wide.

The removal of sea otters has similar effectsas commercial harvest of red sea urchins; thewell known “trophic cascade” of kelpreforestation after release from sea urchingrazing pressure (Sala et al. 1998). Theintense predation effects of sea otters onspecies of commercial invertebrates in theNortheast Pacific including British Columbiaare well known (Watson and Smith 1996).Watson and Smith speculated that theextirpation of sea otters may have allowedinvertebrate stocks to accumulate atunnaturally high levels. However,multispecies relationships within kelp forestecosystems remain poorly understood (Esteset al. 1989; Dewees and Davies 1992).

In Gwaii Haanas, one could reasonablyexpect from sea otter reestablishmentdramatic decreases in red sea urchin and redsea cucumber (Parastichopus californicus)populations and reduction of northernabalone to low numbers of cryptic, crevice-dwellers (Sloan and Breen 1989). In thenorth and east Graham Island areas, therecould be declines in Dungeness crab (Cancermagister) stocks as well. The issue of seaotter-commercial invertebrate speciesinteractions and attendant economic impactswarrant careful consideration as sea ottersare likely to return to Gwaii Haanas naturally(passively) within decades or sooner ifrepatriated by humans.

Active repatriation of sea otters could be afuture sociopolitical issue of importance forGwaii Haanas. Allowance for marineecosystem restoration is established in ParksCanada policy. In the marine component ofthe guiding principles and operational

policies under section 3.1 EcosystemManagement (Parks Canada 1994), thefollowing two subsections are relevant:

• subsection 3.1.2 – “Where marineecosystems or components thereof have beenseriously degraded, Parks Canada willinitiate restoration programs in cooperationwith others”; and

• subsection 3.1.4 – “Extirpated species thatare native to the marine conservation areamay be reintroduced after research hasshown that reintroduction is likely tosucceed and that its probable effects areacceptable within the conservation area andthe surrounding region.”

Pacific Herring Spawn-on-Kelp (SOK)Fisheries and Kelp Mariculture

In part because kelp from the southeast coastof Haida Gwaii is considered to be ofsuperior quality, this is where the firstcommercial (by experimental permit) SOKfishery was initiated in 1971, with a secondexperimental permit issued in 1974 (Shieldset al. 1985). Full commercial productionbegan in 1975 (13 licences) with rapidexpansion, including 11 licences from HaidaGwaii, by 1978. Southeast Haida Gwaii wasalso a location of early development ofmethods for estimating egg deposition as thebasis for herring stock assessment (Haegeleand Schweigert 1985). When available,giant kelp was the herrings’ substrate ofchoice as revealed from SCUBA surveys.

The fishery is based on kelp being deployedin impoundments for deposition of eggs fromgravid herring that have been seined,transported to the impoundment and whichwill later be released after spawning.Mainland harvesters remove kelp mostlyduring February-March from Haida Gwaii.

30

This is done along with the current nine J-licence holders (five of them Haida)permitted to impound herring in HaidaGwaii. There were applications to harvest39 wet tonnes from Haida Gwaii (just northof Skidegate [Lawn Hill] to Cape St. James)in support of the 1999 SOK fishery (Druehl1999 a). Access to the high quality localkelp forests by off-island J-licence holdersis a contentious issue.

From Gwaii Hannas’ view, the major issueis understanding the nearshore ecosystemimpacts of the sanctioned kelp harvest.Although the scale of harvesting is limited,as a matter of management principle, suchunderstanding is warranted for the long-termwell-being of Gwaii Haanas. Moreover, thereturn of sea otters would likely beaccompanied by appreciable kelp forestexpansion as kelp are released from grazingpressure due to sea otter predation on theherbivorous red sea urchins discussed in theabove section. Such expansion could befollowed by SOK industry requests toincrease kelp harvest for theirimpoundments.

An issue requiring verification is thepossible discovery of aberrant giant kelp(perhaps Macrocystis pyrifera) in smallpatches along the southeast coast ofMoresby Island south of Benjamin Point(Druehl and Hopkins 1999). The presenceof M. pyrifera in Gwaii Haanas could haveimplications for the SOK industry and is amost interesting biogeographic question inits own right.

The proposed Marine Conservation AreasAct allows for the possibility of mariculturein Marine Conservation Areas. Given thatGwaii Haanas is a prime collection area ofwild kelp and the need for quality giant kelpfor the SOK industry, there clearly is a

potential demand for a more reliable sourceof kelp through local culture (Druehl 1999a). Kelp culture is more environment-friendly compared, for example, to salmonidculture. There are no concerns aboutescapement, disease introduction, organicenrichment from feed and feces orintroduced chemicals (e.g., antibiotics).Kelp farms require simple floating framesand would appear very similar in size andshape to the k’aaw pens that currently existin Gwaii Haanas. Peak farm activitieswould be in March and April – well beforethe main visitor season starting in June. Keyissues would likely be safety to navigation,visual impact of farms on wildernesssettings and shading effects of farms on theunderlying communities.

Visitor Impacts – Trampling IntertidalSeaweeds

This is the field of “recreation ecology”sensu Liddle (1991), that began with studiesof trampling of terrestrial plants in parks(e.g., Cole and Bayfield 1993) and has sinceexpanded to marine studies on corals andintertidal rocky shores.

Gwaii Haanas had ~ 1,600 visitorscomprising ~10,000-12,000 visitor-day/nights in 1999, which is among the largest“backcountry” (wilderness) visitations in theParks Canada system. Virtually all visitorimpacts on Gwaii Haanas are coastal due tosmall boat access such as kayaks in theintertidal and the immediate back-beach forcamping.

The first Parks Canada initiative onmonitoring visitor impacts on the rockyintertidal was from Pacific Rim NationalPark Reserve (H. Holmes in Rowe et al.1999). The monitoring protocol, based onthe Department of Fisheries and Oceans’

31

“Shorekeeper’s Manual” (Jamieson et al.1999), was initiated in 1997 (Table 7). Datainclude marine plant observations made intide pool, horizontal bench, crevice andvertical rock face substrates. The intertidalkelp monitoring protocol developed forPacific Rim by Druehl and Elliott (1996),that was not focused necessarily on visitorimpact, has not yet been adopted intostandard operations.

There is now a rocky intertidal tramplingliterature in which protected seashore areasin Australia (Keough and Quinn 1998),South Africa (Bally and Griffiths 1993) andthe U.S. (Brosnan 1993; Brosnan andCrumrine 1994) have been the sites ofvisitor impact surveys. The intertidal florafeatures importantly in these studies. Forexample, the rocky intertidal of CabrilloNational Monument, near San Diego, CAhas been monitored by the National ParkService semi-annually since 1990 in whichat least four algae and a seagrass are amongthe target species (Engle and Davis 1996).Generally, foliose algae such as Ulva spp.,Iridaea spp. and fucoid rockweeds aresusceptible to trampling whereas turf-forming species are more resistant. Shiftshave been observed in communitycomposition in favor of low-profile turf overfoliose species. However, Keough andQuinn (1998) caution on the use of intertidalalgae for visitor impact monitors until thenatural variation and patchiness of intertidalspecies are better understood and morediscernable from variation caused by humanimpacts.

Visitor focal points of Gwaii Haanas such asBurnaby Narrows at low tide experienceappreciable visitor traffic includingtrampling the intertidal biota, as articulatedin the Backcountry Management Plan(Gajda 1999). This impact is unquantified

but, given the adoption of the PrecautionaryPrinciple in the proposed Marine ProtectedAreas Act, it could be within Parks Canada’smandate to implement area closure based ontrampling concerns until the science behindsuch intertidal impacts is better understood.The current management recommendation isto promote visitor-vessel float-throughrather than intertidal walking. In fact, theBurnaby Narrows area is closed under theNational Parks Act (i.e., all areas of GwaiiHaanas landward of the high tide line), forvarious non-marine reasons (Gajda 1999).In the longer-term, it would be best toinvestigate trampling in intertidal areas inorder to scientifically back up any long-termarea closure based on marine ecosystemconcerns.

CONCLUSIONS ANDRECOMMENDATIONS

This report provides a baseline marine floraand discusses attendant plant-associatedmanagement issues upon whichconservation enthusiasts, marine biologistsand protected area managers can improvethe marine floristic legacy and itscontribution to nearshore ecosystemstructure and function in Haida Gwaii.

GWAII HAANAS MARINE FLORAKNOWLEDGE

We hope that this report will attract marinebotanists and encourage collection ofvoucher specimens to help complete theregional flora including verifying theabsence of noteworthy species. Appreciabledata gaps occur in all the algal groups foundin Gwaii Haanas, which account for ~49%of regional marine flora biodiversity at thistime. This is all the more important asHaida Gwaii is at the periphery (southernextreme) of the northern seaweed flora. A

32

wide range of botanical opportunities areavailable to investigators including andbeyond the kelp “charismatic megaflora”.

The following recommendations are madebased on this report and the literature:

1) Synoptic surveys for all marine algalphyla should be conducted as knowledge oflocal flora is incomplete for total speciesbiodiversity and for local speciesassemblages.

2) Future surveys should encourageprotocols for establishing species’ absences.

3) Interesting species distributions, such asthe apparent northern limit of the sea palmPostelsia palmaeformis at Hope Island offnorthern Vancouver Island, require furtherinvestigation – is distribution related tohabitat, sea temperature or limits ofpropagule transport (currents) and viability?

4) The genetic distinctiveness of the HaidaGwaii marine flora needs to be assessed andcompared with the region’s mainland florato establish the degree of relatedness.

5) Traditional Haida marine ethnobotanicalknowledge requires further study as it hasbeen relatively little researched beyond plantnaming. Gwaii Haanas is a cooperativemanagement partnership between the HaidaNation and Canada (Hawkes 1996) in whichboth scientific and traditional Haidaknowledge must be seen to contribute tolong-term management.

6) The aberrant Macrocystis species issuewarrants resolution through the simpleretrieval of the (diagnostic) holdfast of theunusually located plants.

7) Parks Canada should establish an

electronic repository providing all interestedparties with access the Gwaii Haanasbiodiversity data sets.

8) The lack of seasonal algal collectionsfrom any location in Haida Gwaii should beattended to.

GWAII HAANAS ECOSYSTEMMANAGEMENT

The vital role of plants to nearshore GwaiiHanaas ecosystem structure and functionand in long-term stewardship initiativesyield the following recommendations:

1) Marine plants such as seaweeds andseagrasses should be used for long-termmonitoring based on physical environment-plant assemblage correlations.

2) Kelp forest monitoring must occur beforenatural (or human) repatriation of sea ottersto enable assessment of post-introductionchanges to these ecosystems.

3) The full geographic extent and an indexof abundance of the invasive species, S.muticum, in Haida Gwaii should beestablished.

4) The relationship of prominent kelp forest-associated species such as sea otters, red seaurchins and northern abalone should beresearched as they impinge on culture,nearshore ecosystem structure and function,and local fisheries.

5) Marine plant values must be includedamong other criteria when implementingzonation, as defined in the proposed MarineConservation Areas Act.

33

LITERATURE CITED

Acheson, S.R. 1995. In the wake of the Iron People: a case for changing settlementstrategies among the Kunghit Haida. Journal of the Royal Anthropological Institute (NewSeries) 1: 273-299.

Adkins, B.D. 1977. A biological and oceanographic analysis of the Queen Charlotte SoundMarine Natural Region. Report prepared for Parks Canada, Hull, P.Q. by Lee and Adkins Ltd., Lantzville, B.C. 103 p.

AMR (Archipelago Marine Research Ltd.). 1997. An environmental assessment of a floatingoperations station in Rose Harbour, Gwaii Haanas. Report prepared for Gwaii HaanasNational Park Reserve, Queen Charlotte, B.C. by Archipelago Marine Research Ltd.,Victoria, B.C. 39 p.

Anderson, P.K. 1995. Competition, predation, and the evolution and extinction of Steller’ssea cow, Hydrodamalis gigas. Marine Mammal Science 11: 391-394.

Anonymous 1998. Marine protected areas - a strategy for Canada’s Pacific coast.Department of Fisheries and Oceans, Pacific Region, Vancouver, B.C. and British ColumbiaLand Use Coordination Office, Victoria, B.C. 27 p.

Augenstein, E.W., D.A. Stow, and A.S. Hope. 1991. Evaluation of SPOT HRV-XS for kelpresource inventories. Photogrammetric Engineering and Remote Sensing 57: 501-509.

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Appendix A

Herbaria staff contacted in the preparation of this report.

Dr. Michael Hawkes, Curator of the Phycological Herbarium, Department of Botany,University of British Columbia, Suite 3529 – 6270 University Boulevard, Vancouver, BC V6T 1Z4 CanadaPhone: (604) 822-5430 e-mail: mhawkes@interchange.ubc.ca

Mr. Michael Shchepanek, Chief Collection Manager of Botany, Canadian Museum of Nature,P.O. Box 3443, Station D, Ottawa, ON K1P 6P4 CanadaPhone: (613) 364-4076 e-mail: mshchepane@mus-nature.ca

Ms. Sarah Gage, Collections Manager, University of Washington Herbarium, Department ofBotany, University of Washington, Box 355325 Seattle, WA 98195-5325 U.S.A.Phone: (206) 543-1682 e-mail: sgage@u.washington.edu

Dr. David Murray, University of Alaska Museum (attn: Herbarium), P.O. Box 756960, Fairbanks, AK 99775 U.S.A.Phone: (907) 474-7108 e-mail: ffdfm@auf.edu

Dr. Paul Silva, Phycology Collection – University Herbarium, University of California –Berkeley, 1001 Valley Life Sciences Building, #2465, Berkeley, CA 94720-2465 U.S.A. Phone: (510) 643-7007 e-mail: psilva@uclink4.berkeley.edu

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Appendix B Part 1

Number of sites at which all marine algae and seagrass have been reported from Haida Gwaiiand Gwaii Haanas National Park Reserve.

Notes.1. ‘-‘ indicates either that the taxa has not been reported for Gwaii Haanas, or that no herbarium specimen(s) for this taxa

are known from Haida Gwaii or Gwaii Haanas.2. Multiple herbarium specimens of the same species were sometimes collected from individual sites.3. UBC = University of British Columbia Phycological Herbarium (2556 specimens).4. CMN = Canadian Museum of Nature herbarium (9 specimens).5. spp. is used where the observations may represent more than one species within a genera.6. These names will be changed according to comb. nov.’s to be published in Gabrielson et al. (in press).

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4 Appendix C

Phylum Chrysophyta (golden algae)Class Sarcinochrysidophyceae

Order SarcinochrysidalesFamily Phaeosacciaceae

Phaeosaccion collinsii 2 1 2 1

Phylum Chlorophyta (green algae)Class Ulvophyceae

Order CladophoralesFamily Cladophoraceae

Chaetomorpha aerea 2 - 2 2Chaetomorpha cannabina 2 - 2 3Cladophora columbiana 8 2 8 4Cladophora hutchinsiae 3 1 3 5Cladophora microcladioides 1 - 1 6Cladophora sericea 2 1 2 7Cladophora spp.5 52 42 3 8Cladophora stimpsonii 2 - 3 9Lola lubrica 1 - 1 10Rhizoclonium riparium 8 - 10 11Rhizoclonium spp. 2 2 - 12Rhizoclonium tortuosum 3 1 3 13

Order CodialesFamily Bryopsidaceae

Bryopsia plumosa 3 - 4 14Derbesia marina 18 5 17 15

Family CodiaceaeCodium fragile 113 71 34 16Codium setchellii 57 37 28 17Codium spp. 4 4 - 18

Order PrasiolalesFamily Prasiolaceae

Prasiola meridionalis 5 2 5 19

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

* Includes ”Schizogonium murale”, a life history stage of Prasiola (Michael Hawkes, personalcommunication).

**Requires confirmation because this species has not been recorded elsewhere on the B.C.coast (M.Hawkes, personal communication).

47

Prasiola spp.* 4 1 1 20Rosenvingiella constricta 1 - 1 21

Order UlotrichalesFamily Acrosiphoniaceae

Acrosiphonia arcta 3 - 2 22Acrosiphonia coalita 13 - 15 | 1 23Acrosiphonia mertensii 12 2 16 24Acrosiphonia saxitilis 5 2 6 25Acrosiphonia spinescens 10 3 14 26Acrosiphonia spp. 30 21 - 27Urospora penicilliformis 2 - 2 28

Family CapsosiphonaceaeCapsosiphon fulvescens 4 - 2 29

Family CollinsiellaceaeCollinsiella tuberculata 6 1 6 30

Family GayraliaceaeGayralia oxysperma 4 - 6 31

Family KornmanniaceaeBlidingia marginata 3 1 3 32Blidingia minima 2 - 3 33Blidingia sp. 1 1 1 34Blidingia subsalsa 3 1 3 35Kornmannia leptoderma 5 1 6 36

Family MonostromataceaeMonostroma grevillei 7 - 6 37Monostroma spp. 3 - 4 38

Family UlotrichaceaeEugomontia sacculata 2 - 3 39Ulothrix flacca 1 - 2 40Ulothrix spp. 5 1 5 41

Family UlvaceaeEnteromorpha clathrata 4 - 4 42Enteromorpha flexuosa 1 - 1 43Enteromorpha intestinalis 20 3 11 44Enteromorpha linza 19 3 15 45Enteromorpha prolifera 4 1 4 46Enteromorpha spp. 47 37 6 47Percursaria percursa 1 - 1 48Ulva crenulata 1** - - 49Ulva fenestrata 31 8 39 | 1 50Ulva spp. 118 91 2 51Ulva taeniata 4 4 - 52Ulvaria obscura var. blyttii 8 1 12 53Ulva/Ulvaria spp. 12 0 - 54

Family UlvellaceaeBolbocoleon piliferum 1 - 1 55Ulvella setchellii 1 - 1 56

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

48

Phylum Phaeophyta (brown algae)Class Phaeophyceae

Order EctocarpalesFamily Acrotrichaceae

Acrothrix sp. 1 - 1 57Family Chordariaceae

Eudesme virescens 5 - 6 58Haplogloia andersonii 8 - 8 59Saundersella simplex 10 1 14 60

Family CoilodesmaceaeCoilodesme bulligera 3 - 3 61Coilodesme californica 2 - 4 62Phaeostrophion irregulare 2 - 2 63

Family DictyosiphonaceaeDictyosiphon foeniculaceus 2 - 2 64

Family EctocarpaceaeEctocarpus parvus 11 2 12 65Ectocarpus siliculosus 7 3 7 66Ectocarpus spp. 2 1 2 67Feldmannia irregularis 1 - 1 68Feldmannia paradoxa var. cylindrica 2 - 2 69Feldmannia sp. 1 - 1 70Hincksia ovata 1 - 1 71Hincksia sandriana 3 - 3 72Hincksia spp. 2 - 2 73Pilayella littoralis 14 2 15 74Pilayella tenella 3 1 1 75Spongonema tomentosum 3 - 4 76

Family ElachistaceaeElachista fucicola 9 3 10 77

Family HeterochordariaceaeAnalipus japonicus 27 15 12 78

Family LeathesiaceaeLeathesia difformis 95 58 41 79

Family MyrionemataceaeHecatonema streblonematoides 1 - 1 80Myrionema globosum 1 - 1 81Myrionema strangulans 3 - 3 82

Family PunctariaceaeMelanosiphon intestinalis 7 1 7 83Punctaria expansa 2 - 2 84Punctaria hesperia 2 - 2 85Soranthera ulvoidea 23 2 30 86

Family RalfsiaceaeRalfsia confusa 1 - 1 87Ralfsia fungiformis 4 - 7 88Ralfsia pacifica 5 - 3 | 1 89Ralfsia spp. 14 14 - 90

Family ScytosiphonaceaeColpomenia peregrina 14 6 16 91Colpomenia spp. 12 10 - 92Petalonia fascia 8 - 5 93Scytosiphon simplicissimus 38 16 27 94

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

49

Scytosiphon spp. 10 1 - 95Family Striariaceae

Stictyosiphon tortilis 2 - 2 96Order Sphacelariales

Family SphacelariaceaeSphacelaria norrisii 1 1 1 97Sphacelaria racemosa 5 1 5 98

Order DictyotalesFamily Dictyotaceae

Dictyota binghamiae 19 6 21 99Order Syringodermatales

Family SyringodermataceaeSyringoderma abyssicola 6 1 6 100

Order DesmarestialesFamily Desmarestiaceae

Desmarestia aculeata 24 8 10 101Desmarestia foliacea 1 1 - 102Desmarestia ligulata 32 12 18 103Desmarestia munda 30 23 5 104Desmarestia viridis 13 4 13 105

Order LaminarialesFamily Alariaceae

Alaria crispa 6 1 7 106Alaria marginata 67 52 11 107Alaria nana 53 41 9 108Alaria praelonga 13 11 3 109Alaria spp. 29 13 7 110Alaria taeniata 1 - - 111Alaria tenuifolia 4 2 3 112Lessoniopsis littoralis 67 50 17 113Pterygophora californica 21 16 3 114

Family LaminariaceaeAgarum clathratum 8 1 8 115Agarum fimbriatum 22 12 2 116Agarum spp. 18 13 2 117Costaria costata 89 58 26 118Cymathere triplicata 45 33 11 119Hedophyllum sessile 116 79 36 120Laminaria bongardiana 106 62 48 121Laminaria complanata 3 - 2 122Laminaria dentigera 8 2 14 123Laminaria ephemera 1 - 1 124Laminaria saccharina 80 45 12 125Laminaria setchellii 114 85 23 126Laminaria spp. 27 2 - 127Laminaria yezoensis 8 - 12 128Pleurophycus gardneri 42 22 22 129

Family LessoniaceaeEgregia menziesii 135 100 33 130Eisenia arborea 3 - 4 131Macrocystis integrifolia 124 84 13 132Nereocystis luetkeana 154 119 24 133

Order Fucales

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

50

Family CystoseiraceaeCystoseira geminata 7 - 11 134

Family FucaceaeFucus gardneri 115 54 49 135Fucus spiralis 4 1 2 136Fucus spp. 94 77 - 137Pelvetiopsis limitata 1 1 - 138

Family SargassaceaeSargassum muticum 14 9 3 139

Phylum Rhodophyta (red algae)Class RhodophyceaeSubclass Bangiophycidae

Order PorphyridialesFamily Porphyridiaceae

Stylonema alsidii 1 - 1 140Order Compsopogonales

Family ErythropeltidaceaeErythrocladia irregularis 3 2 - 141Erythrotrichia carnea 5 1 5 142Erythrotrichia pulvinata 3 1 4 143Porphyropsis coccinea 5 1 5 144Smithora naiadum 15 1 16 145

Order BangialesFamily Bangiaceae

Bangia atropurpurea 11 1 8 146Porphyra abbottae 5 - 6 147Porphyra cuneiformis 3 1 2 148Porphyra fallax 1 - 1 149Porphyra fucicola 3 1 3 150Porphyra gardneri 8 2 9 151Porphyra kanakaensis 1 - 1 152Porphyra lanceolata 1 - - 153Porphyra nereocystis 10 1 8 154Porphyra papenfussii 1 - 1 155Porphyra perforata 24 4 22 156Porphyra pseudolanceolata 3 1 4 157Porphyra schizophylla 5 1 5 158Porphyra smithii 6 1 6 159Porphyra spp. 76 47 29 160Porphyra thuretii 1 - 1 161Porphyra torta 9 3 18 162Porphyra variegeta 1 - 1 163

Subclass FlorideophycidaeOrder Hildenbrandiales

Family HildenbrandiaceaeHildenbrandia occidentalis 6 5 1 164Hildenbrandia rubra 2 - 2 165Hildenbrandia spp. 6 6 - 166

Order AhnfeltialesFamily Ahnfeltiaceae

Ahnfeltia fastigiata 13 3 11 167

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

51

Order CorallinalesFamily Corallinaceae

Bossiella californica ssp. schmittii 15 5 12 168Bossiella chiloensis 6 2 7 169Bossiella cretacea 1 - 1 170Bossiella orbigniana 16 7 12 171Bossiella orbigniana ssp. dichotoma 7 1 8 172Bossiella plumosa 9 1 7 173Bossiella spp. 25 15 -|1 174Calliarthron spp. 9 5 - 175Calliarthron tuberculosum 21 5 21 176Clathromorphum circumscriptum 1 - 1 177Clathromorphum parcum 2 2 2 178Clathromorphum reclinatum 17 4 15 179Clathromorphum sp. 1 - 1 180Corallina frondescens 6 - 6 181Corallina officinalis var. chilensis 31 5 31 182Corallina pilulifera 3 2 1 183Corallina spp. 32 12 - | 1 184Corallina vancouveriensis 81 44 25 185Lithophyllum spp. 7 1 9 186Lithothamnion adeyi6 1 1 1 187Lithothamnion phymatodeum 11 4 5 188Lithothamnion spp. 88 64 - 189Lithothrix aspergillum 1 - 1 190Melobesia mediocris 2 - 1 191Melobesia spp. 2 - - 192Mesophyllum conchatum 2 1 2 193Mesophyllum lamellatum 11 4 8 194Mesophyllum spp. 6 1 6 195Pneophyllum zostericolum 7 - 8 196Pseudolithophyllum muricatum 18 7 14 197Pseudolithophyllum neofarlowii 5 2 3 198Serraticardia macmillanii 15 4 16 199Yamadaia americana 7 2 3 200

Order AcrochaetialesFamily Acrochaetiaceae

Acrochaetium arcuatum 2 1 - 201Acrochaetium densum 2 - 2 202Acrochaetium microscopicum 4 - 4 203Acrochaetium porphyrae 1 - 1 204Audouinella amphiroae6 2 1 2 205Audouinella concrescens6 1 - 1 206Audouinella membranacea 2 - 2 207Audouinella plumosa 1 - 1 208Audouinella plumosa var. variabile6 1 - 1 209Audouinella rhizoidea6 4 - 4 210Audouinella simplex 5 2 - 211Audouinella spp. 4 1 4 212Audouinella thuretii6 2 1 2 213Calaconema daviesii 4 2 4 214Rhodochorton purpureum 5 1 5 215

Order NemalialesFamily Galaxauraceae

Scinaia confusa 12 - 12 216

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

52

Family LiagoraceaeCumagloia andersonii 3 2 - 217Nemalion helminthoides 2 2 - 218

Order PalmarialesFamily Palmariaceae

Halosaccion glandiforme 129 87 36 219Palmaria callophylloides 9 - 12 220Palmaria hecatensis 11 - 16 221Palmaria mollis 22 4 26 222Palmaria spp. 4 - - 223

Family RhodophysemataceaeMeiodiscus spetsbergensis 3 1 2 224Rhodophysema elegans 1 - 1 225Rhodophysema georgii 1 1 1 226

Order CeramialesFamily Ceramiaceae

Antithamnion defectum 18 6 12 227Antithamnion spp. 15 5 - 228Antithamnionella pacifica 2 - 2 229Antithamnionella spirographidis 5 - 3 230Callithamnion acutum 6 - 6 231Callithamnion biseriatum 4 2 3 232Callithamnion pikeanum 42 23 22 233Callithamnion spp. 5 2 4 234Ceramium californicum 4 - 4 235Ceramium codicola 5 2 5 236Ceramium eatonianum 1 - 1 237Ceramium gardneri 4 1 4 238Ceramium pacificum 9 - 7 239Ceramium rubrum 3 1 1 | 1 240Ceramium spp. 16 9 - 241Ceramium strictum 9 1 10 242Ceramium washingtoniense 4 2 4 243Griffithsia pacifica 5 1 6 244Hollenbergia nigricans 2 - 2 245Hollenbergia subulata 6 - 5 246Microcladia borealis 40 22 25 247Microcladia coulteri 4 4 - 248Microcladia spp. 9 2 - 249Neoptilota asplenioides 4 - 7 250Neoptilota californica 5 - 1 251Neoptilota hypnoides 9 6 6 252Pleonosporium vancouverianum 2 1 1 253Pterothamnion pectinatum 4 1 4 254Pterothamnion villosum 7 2 - 255Ptilota filicina 32 6 39 256Ptilota serrata 3 - - 257Ptilota spp. 14 13 - 258Ptilothamnionopsis lejolisea 7 3 9 259Scagelia occidentale 3 - 2 260Tiffaniella snyderae 4 - 4 261

Family DasyaceaeHeterosiphonia densiuscula 15 6 7 262Rhodoptilum plumosum 4 - 4 263

Family Delesseriaceae

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

53

Branchioglossum bipinnatifidum 3 1 1 264Cryptopleura lobulifera 1 1 1 265Cryptopleura ruprechtiana 26 11 17 266Cryptopleura sp. 1 - - 267Delesseria decipiens 15 5 6 268Haraldiophyllum mirabile 2 1 1 269Haraldiophyllum nottii 1 1 1 270Hymenena flabelligera 9 - 8 271Hymenena kylinii 2 - 1 272Hymenena multiloba 6 1 7 273Hymenena setchellii 3 - - 274Hymenena spp. 7 3 1 275Membranoptera dimorpha 6 - 6 276Membranoptera platyphylla 11 4 14 277Myriogramme repens 1 - 1 278Nienburgia andersoniana 2 - 2 279Nitophyllum cincinnatum 12 6 18 280Nitophyllum hollenbergii 1 - 1 281Phycodrys isabellae 1 - 1 282Phycodrys riggii 1 - 1 283Phycodrys setchellii 1 1 1 284Polyneura latissima 39 13 31 285Tokidadendron kurilensis 2 - 3 286

Family RhodomelaceaeAmplisiphonia pacifica 3 - 2 287Herposiphonia plumula 14 1 14 288Neorhodomela larix 104 47 36 | 1 289Neorhodomela oregona 1 - 2 290Odonthalia floccosa 61 6 61 291Odonthalia kamtschatica 3 - 3 292Odonthalia lyallii 5 - 3 293Odonthalia spp. 56 43 3 294Odonthalia washingtoniensis 9 1 6 295Osmundea spectabilis 11 5 6 296Polysiphonia hendryi var. deliquescens 2 1 2 297Polysiphonia hendryi var. gardneri 13 3 16 298Polysiphonia hendryi var. hendryi 12 6 2 299Polysiphonia pacifica 17 5 18 300Polysiphonia pacifica var. disticha 2 - 2 301Polysiphonia pacifica var. gracilis 1 - 1 302Polysiphonia pacifica var. pacifica 3 1 3 303Polysiphonia paniculata 9 2 7 304Polysiphonia spp. 36 24 - 305Polysiphonia tongatensis 4 - 4 306Polysiphonia urceolata 6 2 5 307Pterochondria woodii 9 5 5 308Pterosiphonia bipinnata 23 3 30 309Pterosiphonia dendroidea 22 6 25 310Pterosiphonia gracilis 5 - 3 311Pterosiphonia hamata 11 3 17 312Pterosiphonia sp. 1 - - 313Rhodomela lycopodioides 4 - 5 314

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

* These may be Rhodomela lycopoides or Neorhodomela larix (Michael Hawkes, personalcommunication).

54

Rhodomela spp.* 3 - 1 315Schizochlaenion rhodotrichum 1 - 1 316

Order BonnemaisonialesFamily Bonnemaisoniaceae

Bonnemaisonia geniculata 2 1 3 317Bonnemaisonia nootkana 7 3 4 318Bonnemaisonia sp. 1 1 - 319

Order GelidialesFamily Gelidiaceae

Gelidium coulteri 1 1 - 320Gelidium purpurescens 5 2 4 321Gelidium pusillum 1 - 1 322Gelidium spp. 2 - - 323

Order GigartinalesFamily Choreocolacaceae

Harveyella mirabilis 1 1 - 324Leachiella pacifica 2 - 2 325

Family DumontiaceaeConstantinea simplex 10 4 8 326Constantinea subulifera 4 1 3 327Cryptosiphonia woodii 70 34 18 | 1 328Dilsea californica 4 3 1 329Farlowia mollis 8 1 13 330Neodilsea borealis 2 1 2 331Neodilsea sp. 1 - 2 332Pikea californica 6 2 3 333

Family EndocladiaceaeEndocladia muricata 88 45 41 334Gloiopeltis furcata 61 37 20 335

Family FurcellariaceaeOpuntiella californica 26 11 21 336Turnerella mertensiana 1 - - 337

Family GigartinaceaeChondracanthus corymbiferus 24 18 9 338Chondracanthus exasperatus 18 15 - 339Chondracanthus spp. 35 14 1 340Mazzaella affinis 9 8 1 341Mazzaella californica 5 2 4 342Mazzaella cornucopiae 50 41 7 343Mazzaella heterocarpa 17 6 11 344Mazzaella lineare 11 8 5 345Mazzaella rosea 9 1 10 346Mazzaella splendens 48 26 32 347Mazzaella spp. 22 4 - 348

Family GloiosiphoniaceaeGloiosiphonia capillaris 1 - 1 349

Family KallymeniaceaeCallophyllis crenulata 11 2 9 350Callophyllis edentata 1 1 - 351Callophyllis firma 12 3 6 352

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

* Requires confirmation because this species has not been recorded North of Oregon(M.Hawkes, Personal Communication).

55

Callophyllis flabellulata 16 3 10 353Callophyllis heanophylla 3 1 2 354Callophyllis pinnata 3 1 3 355Callophyllis spp. 15 3 - 356Callophyllis thompsonii 1 - - 357Callophyllis violacea 7 2 5 358Cirrulicarpus n. sp. 1 - 1 359Erythrophyllum delesserioides 12 5 7 360Euthora cristata 3 1 2 361Kallymeniopsis spp. 2 - 2 362Pugetia fragilissima 11 2 13 363

Family PetrocelidaceaeMastocarpus jardinii 13 3 17 364Mastocarpus papillatus 157 99 47 365

Family PeysonneliaceaePeyssonnelia meridionalis 6* 6 - 366Peyssonnelia pacifica 5 2 5 367

Family PhyllophoraceaeAhnfeltiopsis gigartinoides 1 - - 368Ahnfeltiopsis leptophyllus 8 6 2 369Ahnfeltiopsis linearis 4 4 - 370Ahnfeltiopsis spp. 4 3 - 371Stenogramma interrupta 1 - 1 372

Family SchizymeniaceaeSchizymenia pacifica 8 1 5 373

Family SolieriaceaeSarcodiotheca furcata 3 - 3 374Sarcodiotheca gaudichaudii 9 3 6 375

Order HalymenialesFamily Halymeniaceae

Cryptonemia obovata 1 - - 376Cryptonemia spp. 3 1 2 377Grateloupia postelsii 2 - 3 378Grateloupia setchellii 1 - 1 379Halymenia californica 2 - 2 380Halymenia spp. 4 - 6 381Prionitis filiformis 5 2 6 382Prionitis lanceolata 37 23 16 383Prionitis linearis 1 - 1 384Prionitis lyallii 4 2 1 385Prionitis spp. 6 1 2 386

Order RhodymenialesFamily Champiaceae

Gastroclonium subarticulatum 21 10 8 387Family Faucheaceae

Fauchea laciniata 15 4 22 388Fauchea spp. 3 - 1 389

Family Rhodymeniaceae

Herbarium Haida Gwaii1 Specimens1,2 Map No. in Gwaii Haanas UBC3 | CMN4Appendix C

* Family Ruppiaceae is used in place of Potomogetonaceae in The plants of B.C. (Qian andKlinka 1998).

56

Botryocladia pseudodichotoma 4 2 3 390Fryeella gardneri 6 4 5 391Rhodymenia californica 5 2 4 392Rhodymenia pacifica 1 - - 393Sparlingia pertusa 24 1 14 | 1 394

Order GracilarialesFamily Gracilariaceae

Gracilaria pacifica 9 2 4 395Gracilaria spp. 5 3 - 396Gracilariopsis lemaneiformis 8 - 8 397

Order PlocamialesFamily Plocamiaceae

Plocamium cartilagineum 16 15 1 398Plocamium spp. 9 5 - 399Plocamium violaceum 27 12 19 400

Phylum Anthophyta (flowering plants)Class Liliopsida

Order ZosteralesFamily Potogometonaceae*

Ruppia maritima 9 1 - 401Family Zosteraceae

Phyllospadix scouleri 72 44 9 402Phyllospadix spp. 41 34 - 403Phyllospadix torreyi 4 - 4 404Zostera marina 82 45 1 405

57

Appendix B Part 2

Numbers of specimens of marine lichens according to shore zones and substrates in Haida Gwaii.

Notes.1 R=rock/W=drift wood/B=bark of living trees2 preliminary shore zonation categories according to O'Clair et al.(1996)

Shore Zone2

SpeciesSub-

strate1high

intertidal splash saltspray

Amandinea coniops (Wahlenb. in Ach.) M. Choisy ex Scheid. & H. Mayrh. R 1 1 2Arthonia phaeobaea (Norman) Norman R 6 6Aspicilia caesiocinerea (Nyl. ex Malbr.) Arn. R 1 10Aspicilia supertegens Arn. R 1 5 9Bacidia alaskensis (Nyl.) Zahlbr. R 18Bacidina inundata (Fr.) Vezda R 2Buellia griseovirens (Turner & Borrer ex Sm.) Almb. (= B. hassei Imsh., ined.) W 2Caloplaca chlorina (Flotow) H. Olivier R 1Caloplaca citrina (Hoffm.) Th. Fr. R & W 1 5Caloplaca flavogranulosa Arup R 3 2 11Caloplaca inconspecta Arup R 4Caloplaca litoricola Brodo R 8 22Caloplaca rosei Hasse R 3Caloplaca verruculifera (Vainio) Zahlbr. R 1 5 4Catillaria chalybeia (Borrer) Massal. R 1 1 3Catillaria sp. no.2 (corticolous, lge spores) B 1Cliostomum griffithii (Sm.) Coppins B & W 22Coccotrema maritimum Brodo R 1 25Collema fecundum Degel. R 1 2 16Collema flaccidum (Ach.) Ach. R 1Collema furfuraceum (Arn.) Du Rietz var. furfuraceum R 9Diplotomma chlorophaeum (Hepp ex Leighton) Szat. R 2Ephebe lanata (L.) Vainio R 1 16Fuscidea intercincta (Nyl.) Poelt in Poelt & Buschart R 2Fuscidea mollis (Wahlenb.) V. Wirth & Vezda R 3Fuscidea thomsonii Brodo & V. Wirth R 9Hafellia tlellensis Brodo & Sheard W 1Kohlmeyera complicatula (Nyl.) Schatz R 1Lecanora albescens (Hoffm.) Branth & Rostrup (??; see Lecanora sp. no.1) R 1 1Lecanora expallens Ach. W 1 21Lecanora grantii H. Magn. W 1 11Lecanora muralis (Schreber) Rabenh. R 11Lecanora orae-frigidae R. Sant. W 2Lecanora poliophaea (Wahlenb.in Ach.) Ach. s. lat. ("subsp. nobleae Brodo, ssp. nov.") R 2 4Lecanora straminea Wahlenb. ex Ach. R 5Lecanora tenera (Nyl.) Cromb. R 1Lecanora zosterae (Ach.) Nyl. W 5Lecanora sp. no.1 (xanthone-containing, maritime, white, saxicolous; cfr. L. albescens) R 2Lecanora sp. no.5 (xanthone-containing, maritime, saxicolous; cfr. L. contractula) R 1Lecidella effugiens (B. Nilson) Knoph & Hertel in Knoph W 1Lecidella elaeochroma (Ach.) M. Choisy B 11Lecidella scabra (Taylor) Hertel & Leuckert R 2 1Lecidella stigmatea (Ach.) Hertel & Leuckert f. stigmatea R 1 20

Shore Zone2

SpeciesSub-

strate1high

intertidal splash saltspray

58

Ochrolechia subplicans (Nyl.) Brodo subsp. hultenii (Erichs.) Brodo R 2Ochrolechia subplicans (Nyl.) Brodo subsp. subplicans R 2Pannaria laceratula Hue B 9Pannaria leucostictoides Ohlsson B 7Pannaria maritima P.M. Joerg. R 14Pertusaria glaucomela (Tuck.) Nyl. B 30Pertusaria glomerata (Ach.) Schaerer B 1Pertusaria suboculata Brodo & Dibben B 16Physcia caesia (Hoffm.) Füürnr. R 3 30Physcia tenella (Scop.) DC. in Lam. & DC. B 3Polychidium contortum Henssen B 10Polychidium muscicola (Sw.) S.F. Gray R 9Porina pacifica Brodo & R.C. Harris, ined. (n. sp.) R 16Porpidia carlottiana Gowan R 28Punctelia stictica (Duby) Krog R 3Pyrenocollema sublitorale (Leighton) R.C. Harris ex Fletcher in Coppins, P. James & D. Hawksw.

barnacles 5

Pyrrhospora quernea (Dickson) Köörber W 2Ramalina dilacerata (Hoffm.) Hoffm. B 3Ramalina menziesii Taylor B 10Ramalina roesleri (Hochst. ex Schaerer) Hue B 12Rhizocarpon geminatum Köörber R 2 21Rhizocarpon hensseniae Brodo R 14Rhizocarpon hochstetteri (Köörber) Vainio (large spored morph) R 1 30Rhizocarpon hochstetteri (Köörber) Vainio (small-spored morph; = "Rhizocarpon sp. no.1") R 1Rinodina conradii Köörber W 8Rinodina disjuncta Sheard & Tonsberg (Syn. R. columbiensis Sheard, ined.) B 1Rinodina gennarii Bagl. R 2 2Spilonema revertens Nyl. R 1 2 14Tephromela atra (Hudson) Hafellner in Kalb R 3Topelia microspora Brodo, ined. (n. sp.) R 1Tylothallia biformigera P. James & R. Kilias in R. Kilias R 3Verrucaria amphibia Clemente R 1Verrucaria degelii R. Sant. R 2Verrucaria epimaura Brodo R 2 16 1Verrucaria erichsenii Zsch. R 2 1Verrucaria halizoa Leighton R 2Verrucaria maura Wahlenb. in Ach. R 40 40Verrucaria mucosa Wahlenb. in Ach. R 1Verrucaria schofieldii Brodo R 4 8Verrucaria silicicola Fink R 6Verrucaria sp. no.2 (dry rock at edge of beach) R 3Xanthoria candelaria (L.) Th. Fr. R & B 2 31Xylographa hians Tuck. W 13Xylographa opegraphella Nyl. in Rothr. W 3Xylographa vitiligo (Ach.) J.R. Laundon W 7

59

Appendix C

Distribution maps of individual algal and seagrass taxa in Haida Gwaii

Notes.

1. The distribution maps in this appendix are at a scale of almost one to three million(1:3,000,000). The radius of the observation dots at this scale is approximately 500 metres.

2. Two or more dots located close together (< 1000 metres) sometimes may appear as asingle dot on the distribution maps.

60

Chrysophyta and Chlorophyta.

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Map 1 Map 2 Map 3

Map 6Map 5Map 4

Map 7 Map 8 Map 9

Phaeosaccion coll insii Chaetomorpha aerea

Cladophora hutchinsiaeCladophora columbiana

Cladophora sericea Cladophora stimpsonii

Chaetomorpha cannabina

Cladophora microcladioides

Cladophora spp.

61

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Map 15Map 14Map 13

Map 16 Map 17 Map 18

Lola lubrica Rhizoclonium riparium

Bryopsia plumosaRhizoclonium tortuosum

Codium fragile Codium

Rhizoclonium

Derbesia marina

Codium setchellii spp.

spp.

62

Chlorophyta continued.

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Map 24Map 23Map 22

Map 25 Map 26 Map 27

Prasiola meridionalis Prasiola

Acrosiphonia coalitaAcrosiphonia arcta

Acrosiphonia saxiti lis Acrosiphonia

Rosenvingiella constricta

Acrosiphonia mertensii

Acrosiphonia spinescens spp.

spp.

63

Chlorophyta continued.

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Map 33Map 32Map 31

Map 34 Map 35 Map 36

Urospora penicilliformis Capsosiphon fulvescens

Blidingia marginataGayralia oxysperma

Blidingia Kornmannia leptoderma

Collinsiel la tuberculata

Blidingia minima

Blidingia subsalsasp.

64

Chlorophyta continued.

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Map 42Map 41Map 40

Map 43 Map 44 Map 45

Monostroma grevillei Monostroma

UlothrixUlothrix flacca

Enteromorpha flexuosa Enteromorpha linza

Eugomontia sacculata

Enteromorpha clathrata

Enteromorpha intestinalis

spp.

spp.

65

Chlorophyta continued.

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Map 51Map 50Map 49

Map 52 Map 53 Map 54

Enteromorpha prolifera Enteromorpha

Ulva fenestrataUlva crenulata

Ulva taeniata Ulva/Ulvaria

Percursaria percursa

Ulva

var. spp.

spp.

spp.

Ulvaria obs cura blyttii

66

Chlorophyta and Phaeophyta.

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Map 60Map 59Map 58

Map 61 Map 62 Map 63

Bolbocoleon piliferum Ulvella setchellii

Haplogloia andersoniiEudesme virescens

Coilodesme bulligera Phaeostrophion irregulare

Acrothrix

Saundersel la simplex

Coilodesme californica

sp.

67

Phaeophyta continued.

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Map 69Map 68Map 67

Map 70 Map 71 Map 72

Dictyosiphon foeniculaceus Ectocarpus parvus

Feldmannia irregularisEctocarpus

Feldmannia Hincksia sandriana

Ectocarpus siliculosus

Feldmannia paradoxacylindrica

Hincksia ovata

spp.

sp.

var.

68

Phaeophyta continued.

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Map 78Map 77Map 76

Map 79 Map 80 Map 81

Hincksia Pilayella littoralis

Elachista fucicolaSpongonema tomentosum

Leathesia difformis Myrionema globosum

Pilayella tenella

Analipus japonicus

Hecatonema streblonematoides

spp.

69

Phaeophyta continued.

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Map 87Map 86Map 85

Map 88 Map 89 Map 90

Myrionema strangulans Melanosiphon intestinalis

Soranthera ulvoideaPunctaria hesperia

Ralfsia fungiformis Ralfsia

Punctaria expansa

Ralfsia confusa

Ralfs ia pacifica spp.

70

Phaeophyta continued.

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Map 96Map 95Map 94

Map 97 Map 98 Map 99

Colpomenia peregrina Colpomenia

ScytosiphonScytosiphon simplicissimus

Sphacelaria norrisi i Dictyota binghamiae

Petalonia fascia

Stictyosiphon tortilis

Sphacelaria racemosa

spp.

spp.

71

Phaeophyta continued.

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Map 100 Map 101 Map 102

Map 105Map 104Map 103

Map 106 Map 107 Map 108

Syringoderma abyssicola Desmarestia aculeata

Desmarestia mundaDesmarestia ligulata

Alaria crispa Alaria nana

Desmarestia foliacea

Desmarestia viridis

Alaria marginata

72

Phaeophyta continued.

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Map 114Map 113Map 112

Map 115 Map 116 Map 117

Alaria praelonga Alaria

Lessoniopsis li ttoralisAlaria tenuifolia

Agarum clathratum Agarum

Alaria taeniata

Pterygophora californica

Agarum fimbriatum

spp.

spp.

73

Phaeophyta continued.

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Map 118 Map 119 Map 120

Map 123Map 122Map 121

Map 124 Map 125 Map 126

Costaria costata Cymathere triplicata

Laminaria complanataLaminaria bongardiana

Laminaria ephemera Laminaria setchellii

Hedophyllum sessile

Laminaria dentigera

Laminaria saccharina

74

Phaeophyta continued.

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Map 132Map 131Map 130

Map 133 Map 134 Map 135

Laminaria Laminaria yezoensis

Eisenia arboreaEgregia menziesii

Nereocystis luetkeana Fucus gardneri

Pleurophycus gardneri

Macrocystis integrifolia

Cystoseira geminata

spp.

75

Phaeophyta and Rhodophyta.

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Map 141Map 140Map 139

Map 142 Map 143 Map 144

Fucus spiralis Fucus

Stylonema alsidiiSargassum muticum

Erythrotrichia carnea Porphyropsis coccinea

Pelvetiopsis limitata

Erythrocladia irregularis

Erythrotrichia pulvinata

spp.

76

Rhodophyta continued.

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Map 150Map 149Map 148

Map 151 Map 152 Map 153

Smithora naiadum Bangia atropurpurea

Porphyra fallaxPorphyra cuneiformis

Porphyra gardneri Porphyra lanceolata

Porphyra abbottae

Porphyra fucicola

Porphyra kanakaensis

79

Rhodophyta continued.

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Map 177Map 176Map 175

Map 178 Map 179 Map 180

Bossiella orbignianadichotoma Bossiella plumosa

Calliarthron tuberculosumCalliarthron

Clathromorphum parcum Clathromorphum

Bossiella

Clathromorphum circumscriptum

Clathromorphum reclinatum

spp.

sp.

spp.

ssp.

80

Rhodophyta continued.

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Map 181 Map 182 Map 183

Map 186Map 185Map 184

Map 187 Map 188 Map 189

Corallina frondescensCorallina officinalis

chilensis

Corallina vancouveriensisCorallina

Lithothamnion adeyi Lithothamnion

Corallina pilulifera

Lithophyllum

Lithothamnion phymatodeum spp.

spp. spp.

var.

81

Rhodophyta continued.

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Map 195Map 194Map 193

Map 196 Map 197 Map 198

Lithothrix aspergillum Melobesia mediocris

Mesophyllum lamellatumMesophyllum conchatum

Pneophyllum zostericolum Pseudolithophyllum neofarlowii

Melobesia

Mesophyllum

Pseudolithophyllum muricatum

spp.

spp.

82

Rhodophyta continued.

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Map 204Map 203Map 202

Map 205 Map 206 Map 207

Serraticardia macmillanii Yamadaia americana

Acrochaetium microscopicumAcrochaetium densum

Audouinella amphiroae Audouinella membranacea

Acrochaetium arcuatum

Acrochaetium porphyrae

Audouinella concrescens

83

Rhodophyta continued.

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Map 213Map 212Map 211

Map 214 Map 215 Map 216

Audouinella plumosaAudouinella plumosa

variabilis

AudouinellaAudouinella simplex

Calaconema daviesii Scinaia confusa

Audouinella rhizoidea

Audouinella thuretii

Rhodochorton purpureum

spp.

var.

84

Rhodophyta continued.

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Map 222Map 221Map 220

Map 223 Map 224 Map 225

Cumagloia andersonii Nemalion helminthoides

Palmaria hecatensisPalmaria callophylloides

Palmaria Rhodophysema elegans

Halosaccion glandiforme

Palmaria mollis

Meiodiscus spetsbergensisspp.

85

Rhodophyta continued.

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Map 231Map 230Map 229

Map 232 Map 233 Map 234

Rhodophysema georgii Antithamnion defectum

Antithamnionella spirographidisAntithamnionella pacifica

Callithamnion biseriatum Callithamnion

Antithamnion

Callithamnion acutum

Callithamnion pikeanum

spp.

spp.

86

Rhodophyta continued.

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Map 240Map 239Map 238

Map 241 Map 242 Map 243

Ceramium californicum Ceramium codicola

Ceramium pacificumCeramium gardneri

Ceramium Ceramium washingtoniense

Ceramium eatonianum

Ceramium rubrum

Ceramium strictumspp.

87

Rhodophyta continued.

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Map 249Map 248Map 247

Map 250 Map 251 Map 252

Griffithsia pacifica Hollenbergia nigricans

Microcladia coulteriMicrocladia borealis

Neoptilota asplenioides Neoptilota hypnoides

Hollenbergia subulata

Microcladia

Neoptilota californica

spp.

88

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Map 258Map 257Map 256

Map 259 Map 260 Map 261

Pleonosporium vancouverianum Pterothamnion pectinatum

Ptilota serrataPtilota filicina

Ptilothamnionopsis lejolisea Tiffaniella snyderae

Pterothamnion villosum

Ptilota

Scagelia occidentale

spp.

89

Rhodophyta continued.

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Map 267Map 266Map 265

Map 268 Map 269 Map 270

Heterosiphonia densiuscula Rhodoptilum plumosum

Cryptopleura ruprechtianaCryptopleura lobulifera

Delesseria decipiens Haraldiophyllum nottii

Branchioglossum bipinnatifidum

Cryptopleura

Haraldiophyllum mirabile

sp.

90

Rhodophyta continued.

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Map 276Map 275Map 274

Map 277 Map 278 Map 279

Hymenena flabelligera Hymenena kylinii

HymenenaHymenena setchellii

Membranoptera platyphylla Nienburgia andersoniana

Hymenena multiloba

Membranoptera dimorpha

Myriogramme repens

spp.

91

Rhodophyta continued.

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Map 285Map 284Map 283

Map 286 Map 287 Map 288

Nitophyllum cincinnatum Nitophyllum hollenbergii

Phycodrys setchelliiPhycodrys riggii

Tokidadendron kurilensis Herposiphonia plumula

Phycodrys isabellae

Polyneura latissima

Amplisiphonia pacifica

92

Rhodophyta continued.

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Map 289 Map 290 Map 291

Map 294Map 293Map 292

Map 295 Map 296 Map 297

Neorhodomela larix Neorhodomela oregona

Odonthalia lyalliiOdonthalia kamtschatica

Odonthalia washingtoniensisPolysiphonia hendryi

deliquescens

Odonthalia floccosa

Odonthalia

Osmundea spectabilis var.

spp.

93

Rhodophyta continued.

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Map 303Map 302Map 301

Map 304 Map 305 Map 306

Polysiphonia hendryigardneri

Polysiphonia hendryihendryi

Polysiphonia pacificagracilis

Polysiphonia pacificadisticha

Polysiphonia paniculata Polysiphonia tongatensis

Polysiphonia pacifica

Polysiphonia pacificapacifica

Polysiphonia spp.

var. var.

var.var. var.

94

Rhodophyta continued.

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Map 312Map 311Map 310

Map 313 Map 314 Map 315

Polysiphonia urceolata Pterochondria woodii

Pterosiphonia gracilisPterosiphonia dendroidea

Pterosiphonia Rhodomela

Pterosiphonia bipinnata

Pterosiphonia hamata

Rhodomela lycopodioidessp. spp.

95

Rhodophyta continued.

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Map 321Map 320Map 319

Map 322 Map 323 Map 324

Schizochlaenion rhodotrichum Bonnemaisonia geniculata

Gelidium coulteriBonnemaisonia

Gelidium pusillum Harveyella mirabilis

Bonnemaisonia nootkana

Gelidium purpurescens

Gelidium spp.

sp.

96

Rhodophyta continued.

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Map 330Map 329Map 328

Map 331 Map 332 Map 333

Leachiella pacifica Constantinea simplex

Dilsea californicaCryptosiphonia woodii

Neodilsea borealis Pikea californica

Constantinea subulifera

Farlowia mollis

Neodilsea sp.

97

Rhodophyta continued.

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Map 339Map 338Map 337

Map 340 Map 341 Map 342

Endocladia muricata Gloiopeltis furcata

Chondracanthus corymbiferusTurnerella mertensiana

Chondracanthus Mazzaella californica

Opuntiella californica

Chondracanthus exasperatus

Mazzaella affinisspp.

98

Rhodophyta continued.

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Map 348Map 347Map 346

Map 349 Map 350 Map 351

Mazzaella cornucopiae Mazzaella heterocarpa

Mazzaella splendensMazzaella rosea

Gloiosiphonia capillaris Callophyllis edentata

Mazzaella lineare

Mazzaella

Callophyllis crenulata

spp.

99

Rhodophyta continued.

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Map 352 Map 353 Map 354

Map 357Map 356Map 355

Map 358 Map 359 Map 360

Callophyllis firma Callophyllis flabellulata

CallophyllisCallophyllis pinnata

Callophyllis violacea Erythrophyllum delesserioides

Callophyllis heanophylla

Callophyllis thompsonii

Cirrulicarpus n. sp.

spp.

100

Rhodophyta continued.

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Map 361 Map 362 Map 363

Map 366Map 365Map 364

Map 367 Map 368 Map 369

Euthora cristata Kallymeniopsis

Mastocarpus papillatusMastocarpus jardinii

Peyssonnelia pacifica Ahnfeltiopsis leptophyllus

Pugetia fragilissima

Peyssonnelia meridionalis

Ahnfeltiopsis gigartinoides

spp.

101

Rhodophyta continued.

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Map 375Map 374Map 373

Map 376 Map 377 Map 378

Ahnfeltiopsis linearis Ahnfeltiopsis

Sarcodiotheca furcataSchizymenia pacifica

Cryptonemia obovata Grateloupia postelsii

Stenogramma interrupta

Sarcodiotheca gaudichaudii

Cryptonemia spp.

spp.

102

Rhodophyta continued.

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Map 384Map 383Map 382

Map 385 Map 386 Map 387

Grateloupia setchellii Halymenia californica

Prionitis lanceolataPrionitis filiformis

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Halymenia

Prionitis linearis

Prionitis spp.

spp.

103

Rhodophyta continued.

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Map 393Map 392Map 391

Map 394 Map 395 Map 396

Fauchea laciniata Fauchea

Rhodymenia californicaFryeella gardneri

Sparlingia pertusa

Botryocladia pseudodichotoma

Rhodymenia pacifica

Gracilaria pacifica spp.Gracilaria

spp.

104

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Map 402Map 401Map 400

Map 403 Map 404

Gracilariopsis lemanaiformis Plocamium cartilagineum

Ruppia maritimaPlocamium violaceum

Phyllospadix

Plocamium

Phyllospadix scouleri

Phyllospadix torreyispp.

spp.

Map 405

Zostera marina