The Pacific coast of North America 199

19 The of THE PACIFIC COAST OF NORTH AMERICA

S. Wyllie-Echeverria J.D. Ackerman

he region along the Pacific coast of North America In contrast, soft-bottom habitats in the subtidal extending from the Baja Peninsula in Mexico and intertidal zones and estuaries are more commonly Tthrough Alaska includes a wide variety of associated with in the genus Zostera, which can ecosystems ranging from subtropical through arctic in form large monotypic stands in the Northeast Pacific a northerly transect. Given the nature of the leading estuaries, and mixed stand populations of Zostera edge coast and the resultant paucity of large regions marina and Zostera japonica, and sometimes where soft sediments can accumulate, one would maritima, in estuaries from southern British Columbia, expect a rather limited diversity of marine angiosperms Canada, to Coos Bay, Oreg~n".~'.Zostera marina or seagrasses. However, a reasonably large number of provides important habitat for migrating waterfowl, species exist in this region for a number of reasons, juvenile salmon, resident forage fish, invertebrates and related in part to the ability of members of the genus wading birds'", and Zostera japonica is commonly eaten Phyllospadix to colonize rocky shores. Eight by resident and migratory waterf~wl"~'.Noteworthy is species are recognized: Halodule wrightii, Ruppia the use of Zostera marina as substrate for the laying of maritima, Zostera marina, Zostera japonica, Zostera Pacific herring IClupea harengus pallasil roe; the roe is asiatica, , also used by humans"'. and Phyllospadix torreyjl-'I. Four of them, Zostera Whereas little is known about the primary marina, Phyllospadix scouleri, Phyllospadix serrulatus production rates for Zostera japonica, Zostera marina and , have probably been growing in productivity can be quite high on an annual basis 184- the region since the PlioceneIa; one, Zostera japonica, 480 g carbon/m2/yearl and standing stocks may cover is a recent addition to the northeast Pacific flora, being many hectares of ~eafloor'~."'.For example, the large introduced as a result of oyster enhancement populations at lzembek Lagoon, Alaska. United States program^'^'; and little is known about the phyto- 1160 ha) and Laguna Ojo de Liebre, Baja California, geographic history of the three other species (Zostera Mexico I175 ha), which are the primary staging asiatica, Ruppia maritima and Halodule wrightiil. grounds for migratory waterfowl, may be the largest In terms of ecosystems, members of the genus Zostera marina ecosystems in the ~orld"~~~~'.In Phyllospadix Ithe surfgrassesl dominate the rocky addition, pre-contact First Nations peoples recognized subtidal and intertidal zones, where their condensed Zostera marina and its ecosystems as valuable rhizomes allow them to colonize hard substrates. The cultural and food resources. In British Columbia a three species in the genus Phyllospadix IPhyllospadix number of First Nations people (Nuu-chah-nulth, serrulatus, Phyllospadix scouleri and Phyllospadix Haida and Kwakwaka'wakwl ate fresh rhizomes and torrey~lare endemic to the Northeast Pacific12'. Both leaf bases or dried them into cakes for winter food"3'. Phyllospadix torreyi and Phyllospadix scouleri were Moreover, the Seri Indians living on the Gulf of widely used in the region by indigenous people before California in Sonora, Mexico, used the Zostera marina European contact"'. For example the flowers of seeds to make flour"". Phyllospadix torreyi were sucked for sweetness by Ruppia maritima grows in many of the brackish children of the Makah people who live on the Olympic water coastal lagoons from Alaska south to Me~ico".'~'. Peninsula of Washington state, and leaves of the same Interestingly, Ruppia maritima was recognized as a were woven into pouches by the coastal Chumash separate species lfrom Zostera marina1 by the Seri in the Channel Islands of California. elders but was not used by them"". M 200 WORLD ATLAS OF SEAGRASSES

The last two species, namely Zostera asiatica BIOGEOGRAPHY and Halodule wrightii, have rarely been the focus of Zostera marina [or eelgrass) is the dominant species in biogeographic investigation with~n this region. terms of biomass and habitats on the Pacific coast of Zostera asiatica was found recently at three sites North America, where it grows in: in southern California"' and, Leaving aside some o the shallow waters of the continental shelf; regional studies documenting the presence of o the Gulf of California [Sea of Cortezl; Halodule wrightii in the Gulf of Cal~fornia'",there are o coastal lagoons such as San Quintin, Baja no studies that discuss the habitat value or California, Mexico, and lzembek Lagoon, ~laska""; autecology of these plants in the Northeast Pacific n estuaries formed by tectonic processes like San region. Francisco Bay;

forage on seagrass flats (primarily Zostera marina] non-stop, three-day transoceanic flight to Zostem ; from Alaska to Mexico. In Late August, afer raising marina and Ruppia maritma beds in Baja California 7 young in theYukon-Kuskokw~mDelta 16I0N, 165OW1, at Bahia San Quintin I3O0N. 116"W; see photograph -1 flocks gather at lzembek Lagoon [55"N, 163"W) to below right), Laguna Ojo de Liebre [27"N,114"v\ll graze on one of the largest intertidal stands of and Laguna San lgnacio [26"N, 113°W1. Zostera marina in the world [see photograph below Spring migration coincides with midday

locations like Morro Bay and Humbotdt Bay, Cabfornia; South Slough and Yaquina Bay, Oregon; Willapa Bay and Padilla Bay, Washington: and : Boundary Bay. British Columbia, Canada. International conservation efforts by the United States, Canada and Mexico are under way at - winter~ngand migration stopover sites along the) eastern Pacific Flyway to pmtect seagrass habifats - In coastal embayments and estuaries. ----

In collaborat~onw~th Dav~d Ward (US Geological Survey, Anchorage] - and Dr S~lvlaIbarra-Obando ICentm de InvestigacionesCientiitcayde - Educacldn Superlor de Ensenada. Baja California, Mexical. - 4

4. .%f' u coastal fjords similar to Puget Sound, <.. f, 160' W 150' W ' .' .-. Washington"'...... 4 *'. , 8 ERIN G Yu~uII . I' V.!/U>. . .. It is found along the coast of British Columbia SEA \: including the coasts of Vancouver Island and Queen i: , USA Charlotte Islands (Haida Gwaii) in sheltered bays and , . .ri3-.... : Alaska :, ' -> < coves including Bamfield Harbour and Sooke Basin'". , . - - The species also extends well into Alaskan waters to the i. .?_ ..1. ;: Arctic Circle115'.In the intertidal zone Zostera marrna can . , co-mingle with Zostera japonica in the Pacific Northwest fi. and Ruppia maritima in Baja Calif~rnia",'~'. Whereas the majority of Zostera marina populations are perennial, annual populations 1e.g. Bahia Kino, Gulf of California, Mexico; Yaquina Bay, Oregon, United States] in which 100 percent of the -po@ah+i~e +eeer&ug sb&lhat m.Lfrom- seeds each year have been reported".'*'. The appearance of branched reproductive shoots, a dimorphic expression quite distinct from the ribbon- shaped leaves of the vegetative shoots, begins to occur as water temperatures warm in the spring. In the Northeast Pacific, reproductive shoots are visible in February at southern sites such as Baja California, Mexico and southern California; in late March or early April in Puget Sound, Washington; and as late as June in northern sites Like lzembek Lagoon. Alaska. Flowering phenology is protogynous and the emergence of stigmas and then anthers effect the release, transport and capture of pollen, which rotate in the shear around stigmas'"'. Zostera marina is monoecious; however the release of pollen and its stigmatic capture is separated in time to promote an outcrossing breeding stern''^^'^'. !hhoBay In a region-wide analysis of population Channel. - i.., ~tructure''~',Alberte and colleagues found that: Islands san we& i-.. ,;, o there was high genetic diversity among Zostera ...... marina populations in the region; -. San Quintin : MEXICO ' . - o gene flow restriction existed for populations that som < : .t+ohh were near each other; KiM L*,gunu (80~e ~icl~re ---...-++ . o intertidal plants in disturbed environments were .... I~I~III~II~ILtul.2 ,' .' less~verssugm~thmrtlror~8is~-- sites.

In a subsequent study, focused on San Diego, Map 19.1 California, and Baja California, Mexico, Williams and The Pacific coast of North America Davis discovered that transplanted Zostera marina populations were less diverse genetically than naturally the early part of the 1900~~~'.At several sites from occurring population^'^^'. southern British Columbia, Canada, to southern Although Zostera japonica is typically smaller Oregon, Zostera japonica co-mingles with Zostera than Zostera marina, it can be confused with the marina [and occasionally Ruppia maritimb9'l in the intertidal growing habit of Zostera marina Ivar. intertidal region of many estuaries1''. Zostera japonica typicali2". However Zostera japonica commonly grows is restricted in its northerly extent to the region near higher in the intertidal zone and has an open (as the city of Vancouver including Boundary Bay and opposed to tubular) leaf sheath characteristic of its Tsawwassen, where it is found primarily in the upper subgenus Z~sterella'~'.It is a possible invader to the intertidal zones in muddy or silty areas'". Some region coming by way of the oyster trade with Japan in unconfirmed reports also exist of the species further 202 WORLD ATLAS OF SEAGRASSES

ZOSTERA MARINA [TS'ATS'AYEM~AND NCOUVER ISLAND, CANADA

.- 2 Chief Adam Dick [Kwaxsistala] and Kim Recalma- -= 7 Clutesi [OqwiloGwal are members of the -E Kwakwaka'wakw Nation on the northeast coast of P LL Vancouver Island, British Columbia, Canada. Both 2 are keenly aware of the value of Zostera marina or -0 20 ts'dts'ayem from oral tradition of their nation. They recall that at Grassy Point or wiwasalth, ts'dtsbyem is collected with a Long thin pole or k'elpawi that is stuck into the substrate, rotated to entwine the leaves of ts'dts'ayern, and pulled from the bottom to reveal leaves, rhizomes and roots. On removal the plants are peeled exposing the tender soft tissue of the leaf base. The leaves are then wrapped around the rhizome, dipped in klina [eulachon IThaleichthyspacificusI grease1 and eaten as a ceremonial food. Whereas Grassy Point has both cultural and ecological value, Chief Dick, Kim Recalma-Clutesi and others of the Kwakwaka'wakw Nation have concern about regional and global practices that threaten the survival of ts'its'ayem.

Chief Adam Dick twisting the s 35s [Zostera rn, dipping in eulachon [Thaleichthys pacificusl grease, Plants were In collaboration with Chief Adam Dick and Kim Recalrna-CLutesi harvested at Deep Bay on the east coast of Vancouver Island on IKwakwaka'wakw Nation1 and Dr Nancy J. Turner [University of 28 July 2002. Victoria. Victoria. Canada].

- ,= -

north in the Strait of Georgia"'. Large stands are and Phillips and ~enez' describe the habitat and present in Boundary Bay, southern British Columbia, regional distribution of the three species. Phyllospadix Canada, and Padilla Bay and Willapa Bay, serrulatus grows in the upper intertidal zone l+1.5 m to Washington""'. The sediments and fauna within Zostera mean lower low water1 on the outer coasts of Alaska, japonica beds were found to be largely similar to those British Columbia, Washington and Oregon. Its found in Zostera marina beds in Oregon, although distribution is often confused with Phyllospadix some differences in sediment grain size and organic scouleri, which inhabits the lower intertidal and shallow constituents were observed'22'. Zostera japonica has subtidal zone. It can be locally quite common, as on been shown to be important to resident and migratory Graham Island (Haida Gwaiil"', and has a distribution waterfowl in Boundary Bay""', and is used as habitat by that extends from southeast Alaska to Baja California, epibenthic crustaceans in Padilla Bay. To the best of our although it is reported to be more abundant north of knowledge, there are no other studies Linking Zostera Monterey, Calif~rnia'~.'I. Phyllospadix torreyi grows at japonica to secondary consumers either as a food or greater depths and is generally more abundant on the habitat in North America. exposed parts of the coast and even in tidal pools with The three species of the genus Phyllospadix are sandy bottoms, which are typically devoid of the other found on exposed rocky coasts in the surf zone and in two Phyllospadix species. The distribution of tide pools in the intertidal zone where their condensed Phyllospadix torreyi nearly overlaps Phyllospadix rhizome allows them to attach to hard substrates. Three scouleri, but it is more abundant south of Monterey, of the five species in this North Pacific genus are found California. The Lack of information on its distribution on the west coast of North America. Turner and L~cas'~~'may be related to the difficulty of making collections in P.. -C(% d- Y ,: I :. ..' The Pacific coast of North America 203

energetic habitats where Phyllospadix torreyi is found'". maritima occur in coastal lagoons and estuaries Whereas little is known about the biogeography of these throughout British Columbia including the Haida species, studies have revealed aspects of their Gwaii'". Further south Ruppia maritima occurs at many autecology and life history such as the adaptations of sites influenced by saline water in Washington, Oregon, seeds and roots to cling to surfaces in the rocky California and northern Mexi~o"~'~'.The leaves, intertidal zone123."I. Phyllospadix spp. form patches of rhizomes and seeds of this plant are eaten by resident various sizes in the surf zones, except Phyllospadix and migratory waterfowl. serrulatus, which is often found in more protected Halodule wrightii is a subtropical species that en~ironments'~'.Plants in the genus Phyllospadix are occurs in the Gulf of California off the coast of largely clonal, and can be of a single sex due to the mainland Mexico['.2', and Zostera asiatica is known to dioecious nature of the genus in this region. occur in three subtidal regions in central California. Few studies have documented the habitat value of United States, where it forms underwater forests Phyllospadix; however, infaunal polychaetes are known ca 3 m tallI3'. More work is necessary to elucidate the to live in the rhizome mats of Phyllospadix scouleriand life history traits and habitat value of these species in Phyllospadix torreyi. Surfgrass wrack, identified as this region. Phyllospadix torreyi, has also been found in the macrophyte detritus layers in submarine canyons, in HISTORICAL PERSPECTIVES southern and central Calif~rnia'~~'.This decomposing Potential changes in the standing crop and areal vegetation provides food and habitat for deep-sea extent of Zostera marina have concerned natural benthic fauna125'.In terms of commercially important resource managers in the Northeast Pacific for more species, researchers in southern California found that than two decades1&26'. This concern is primarily a in their larval pelagic stage, spiny lobsters, Panulirus function of the habitat value provided by the large interruptus, were attracted to experimental treatments ecosystems created by these plants. Changes in the containing Phyllospadix torreyi. distribution of Zostera japonica have received less Ruppia maritima is a variable plant with a number attention. This species is an exotic but provides of named varieties with characteristic features, and is valuable waterfowl habitat"". Given this, and the fact found in both freshwater and marine habitats. Ruppia that Zostera japonica has not yet been shown to maritima var. spiralis occurs along the southern coast negatively impact the indigenous Zostera marina, of Alaska including the Alaska Peninsula, in British prompts some to argue for detailed resource Columbia and in California"'. Varieties longipes and inventories. Information about changes in the local or

Case Study 19.3 THE LINK BETWEEN SEAGRASSES AND HUMANS IN PICNIC COVE, SHAW ISLAND, WASHINGTON, UNITED STATES

Picnic Cove is a sheltered embayment on the .-trn southeast corner of Shaw Island, which is centrally 0 located in the San Juan Archipelago in the Pacific .-$a Northwest [see photograph; 48'35'N 122'57'W). P ui It contains a Zostera marina meadow of iri ca 0.05 km2'"' and a very small patch of Zostera -o E japonica, in addition to multi-layered shell middens on the low bank at the head of the cove, which indicate historical use by coastal Salish people. After European contact, Picnic Cove became a favorite picnic spot. It is now the site of a Long-term monitoring station for Washington State Department of Natural Resources' Submerged Vegetation Monitoring he San Juan Archipelago with Project, and is the location of quadrat-based insert of Picnic Cove on Shaw investigations by S. Wyllie-Echeverria. University of Island, Washington, United Washington. States. 204 WORLD ATLAS OF SEAG.RASSES d

However, there is anecdotal information to suggest that Table 19.1 losses of Zostera marina have occurred in the two Zostera marina and Zostera japonica basal area cover in the largest estuaries on the west coast of the continental Northeast Pacific United States - Puget Sound and San Francisco Bay'n' - and we suspect losses have occurred at other sites as Country Region Area lkm21 well. Widespread efforts to monitor and map this

USA Port Clarence, AK'28s291 4.2 species in this region should help to determine if local USA Safety Lagoon, AK'". 291 9.1 and regional losses continue in the 21st century. More USA lzernbek Lagoon, AK'~~' 159.5 effort is needed to develop a programmatic response USA Kinzarof Lagoon, AK"~.291 8.7 for comprehensive resource inventories of the other USA East Prince William Sound, AK'~" 4.4 seven seagrass species[26'. Canada Roberts Bank, BC'~" 4' Canada Boundary Bay, BC"" 56' SEAGRASSCOVERAGE USA Puget Sound, WA'~~' 200 Whereas an estimate of the seagrass cover is Sites within Puget Sound, WA marginally possible for Zostera marina, it is not Padilla ~ay1~~' 32' possible for the other seven species. The conservative King ~ounty'35' 2 estimate for Zostera marina is approximately 1 000 km2 USA Grays Harbor, WA"' 47.3 and is based on studies cited in Table 19.1 and our USA Willapa Bay, WA'~~' 159' personal knowledge of sites not yet mapped. USA Netarts Bay, OR'"' 3.4 USA Yaquina Bay, 0~'~~' 0.9' THREATS USA Tillamook Bay. OR'39' 3.6 The following discussion of the present and potential USA Coos Bay, OR'^' 0.01' threats to seagrasses is based on some information USA Hurnboldt Bay, CA'~' 12.2 documented by Phillips"', Wyllie-Echeverria and USA Tomales ~af' 3.9 and studies therein, as well as a degree of USA San Francisco ~ay'~~' 1.9 unpublished observation and conjecture. USA San Diego ~ayl'3' 4.4 Coastal modifications and overwater structures in Mexico Bahia San ~uintin'~" 20 the form of ferry terminals, commercial docks, and Mexico Laguna Ojo de ~iebre'~" 175 smaller residential docks and floats threaten the Mexico Laguna San ~~nacio'~~' 53 survival of species that could be shaded in either soft- bottom or rocky littoral zones. Shoreline armoring, Note: * Includes both Zostera marina and Zostera japonica. which can alter the trajectory of reflected wave energy, may also displace seagrasses. The direct removal of Source: Various sources - see individual references by regions. seagrasses through maintenance dredging is a rare occurrence in the Northeast Pacific, but resuspension of sediment associated with activity outside the regional abundance of the other seagrass species in seagrass zone may reduce transmission of light and/or the Northeast Pacific remains largely unkn~wn"~'. bury plant populations. The deposition of upland soils Direct use of seagrasses by humans for food, into the littoral zone as a result of industrial, com- technology and medicine in the Northeast Pacific was mercial and residential development may smother widespread before European c~ntact'~~'~~"'.However, and/or kill seagrass. Moreover, modifications to the use now is quite localized and involves the weaving of coastline projection may alter longshore current Phyllospadix spp. as a decorative element in small patterns resulting in changes to water clarity. personal baskets and the collection of Zostera marina Recreational watercraft (powerboats, jet skis, plants for green mulch or the protection of culturally etc.1 may scar seagrasses in soft-bottom environments significant sites (see Case Study 19.21. The United resulting in the fragmentation of populations and the States Department of Agriculture investigated the subsequent loss of wildlife habitat. Whereas larger potential use of Zostera marina as a cultivar in coastal vessels (ferries, freighters, tankers, etc.1 rarely venture desert ecosystems during the 1980% but we are into shallow waters, accelerated currents associated unaware of any projects to further this goal. with propeller wash connected with landing and getting It is difficult to ascertain the extent of seagrass under way may displace seagrasses and affect current losses due to coastal development and population flow during pollen and seed release. The swing of expansion since the beginning of the 20th century, as anchor chains, and chains and lines connected to no baseline data exist prior to the onset of these permanent buoys, can uproot plants and leave Any attempt to do so would be conjecture. permanent scars in populations. The Pacific coast of North America 205

Rack and rope culture techniques used in commercial shellfish culture may shade the bottom and alter nutrient regimes and current flow, which may result in the loss of seagrass cover in localized areas. Human trampling associated with the harvest of market-sized oysters from stakes used to set and grow oyster spat can also result in reductions in seagrass cover. Moreover, recreational clam removal using shovels can destroy meters of seagrass cover. Spills associated with oil production from offshore oil platforms such as those located on the continental shelf of southern California or the transport by oceanic tankers from Alaska to southern ports in Washington and California may result in the death of seagrasses in the littoral zone depending on the intensity and duration of the spill. Proposals for Small net bag or flexible basket woven from the leaves of offshore oil development in Canada, the United States Phyllospad~xtorreyi, found at Santa Rosa Island, Californ~a,and and Mexico are most problematic in this regard. dated 1100-1500 In the Common Era. Episodic events such as ENS0 IEl Niiio Southern Oscillation1 and interdecadinal variation have the potential to alter ocean temperature and rainfall regimes, which may affect local populations and may also operate on the regional scale. However, a preliminary investigation in Puget Sound found that both biomass and productivity of a subtidal Zostera marina population increased during an El Niiio year 11991-921 demonstrating the need for time-series data collection to evaluate the status and trend of seagrass ecosystems. Subduction associated with the nearshore plate tectonic activity may alter the shape and size of the littoral zone, reducing or eliminating the seagrass cover, as in the case of the 1964 Alaska earthquake. The fragmentation of populations caused by natural or anthropogenic disturbance can also provide habitat for introduced species such as the mussel, Musculista senhousia, which can in turn prevent regrowth into fragmented areas, potentially leading to a more widespread decline in seagrass cover.

POLICY OPTIONS AND SEAGRASS PROTECTION It is not clear that federal, provincial or state, or local Zostera marina prairie adjacent to the ferry terminal on Whidbey administrative laws and ordinances recognize the Island In central Puget Sound. Prairie density is Influenced by both eight seagrass species in the Northeast Pacific. the overwater structure and ferry pmpwash However, in the United States, Canada and Mexico, protection is afforded to Zostera marina because the ecosystems provided by this plant are valuable habitat AUTHORS for commercially and recreationally important species Sandy Wyllie-Echeverria, School of Marine Affairs, Box 355685, such as Pacific salmon (Oncorhynchus spp.1, Pacific University of Washington, Seattle, WA, USA, 98105-6715. herring IClupea harengus pallas11 and black brant Tel: t1 360 468 6619; 293 0939. Fax: +I 206 543 1617. E-mail: [Branta bernicla nigricansIl8'. In Washington state, [email protected] Zostera japonica is also protected, but to the best of our knowledge no other seagrasses are protected by Josef Daniel Ackerman. Physical Ecology Laboratory, Univers~tyof administrative code in the Northeast Pacific. Northern British Columbia, Prince George, BC, Canada, V2N 429. 206 WORLD ATLAS OF SEAGRASSES

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