Vol. 32, No. 1 January 2004 FREMONTIAvolume 32:1, january 2004 a journal of the california native...

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

A JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

FREMONTIA

$5.00 (Free to Members)

Vol. 32, No. 1 January 2004

IN THIS ISSUE:IN THIS ISSUE:IN THIS ISSUE:IN THIS ISSUE:IN THIS ISSUE:

SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES by Paul C. Silva 3 3 3 3 3

CALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDS by Kathy Ann Miller 1010101010

CALIFORNIA SEAWEED INDUSTRY CALIFORNIA SEAWEED INDUSTRY CALIFORNIA SEAWEED INDUSTRY CALIFORNIA SEAWEED INDUSTRY CALIFORNIA SEAWEED INDUSTRY by Paul C. Silva 1616161616

DISTRIBUTION OF CALIFORNIA SEAWEEDS DISTRIBUTION OF CALIFORNIA SEAWEEDS DISTRIBUTION OF CALIFORNIA SEAWEEDS DISTRIBUTION OF CALIFORNIA SEAWEEDS DISTRIBUTION OF CALIFORNIA SEAWEEDS by Paul C. Silva 1818181818

INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS by Kathleen Dickey 2727272727

SPECIAL ISSUE: SEAWEEDSSPECIAL ISSUE: SEAWEEDSSPECIAL ISSUE: SEAWEEDSSPECIAL ISSUE: SEAWEEDSSPECIAL ISSUE: SEAWEEDS

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

The California Native Plant Society(CNPS) is a statewide nonprofit orga-nization dedicated to increasing theunderstanding and appreciation ofCalifornia’s native plants, and to pre-serving them and their natural habi-tats for future generations.

CNPS carries out its missionthrough science, conservation advo-cacy, education, and horticulture atthe local, state, and federal levels. Itmonitors rare and endangered plantsand habitats; acts to save endangeredareas through publicity, persuasion,and on occasion, legal action; pro-vides expert testimony to governmentbodies; supports the establishment ofnative plant preserves; sponsors work-days to remove invasive plants; andoffers a range of educational activitiesincluding speaker programs, field trips,native plant sales, horticultural work-shops, and demonstration gardens.

Since its founding in 1965, the tra-ditional strength of CNPS has beenits dedicated volunteers. CNPS ac-tivities are organized at the local chap-ter level where members’ varied in-terests influence what is done. Volun-teers from the 33 CNPS chapters an-nually contribute in excess of 87,000hours (equivalent to 42 full-time em-ployees).

CNPS membership is open to all.Members receive the quarterly jour-nal, Fremontia, the quarterly statewideBulletin, and newsletters from theirlocal CNPS chapter.

Fremontia logo (by L.A. Vorobik) re-printed from The Jepson Manual, J.Hickman, Ed., 1993, with permissionfromthe Jepson Herbarium, UC. © Re-gents of the University of California.

CALIFORNIA NCALIFORNIA NCALIFORNIA NCALIFORNIA NCALIFORNIA NAAAAATIVETIVETIVETIVETIVE

PLANT SOCIETYPLANT SOCIETYPLANT SOCIETYPLANT SOCIETYPLANT SOCIETY

Dedicated to the Preservation ofthe California Native Flora

VOL. 32, NO. 1, JANUARY 2004

FREMONTIAFREMONTIAFREMONTIAFREMONTIAFREMONTIA

Copyright © 2004California Native Plant Society

STAFFSacramento Office:Executive Director . . . . Pamela C.

Muick, PhDDevelopment Director . . . Michael

TomlinsonMembership Coordinator . . Marin

LemieuxSales Manager . . . . . . . Paul MaasBookkeeper . . . . Lois Cunningham

At Large:Fremontia Editor . . . . . Linda Ann

Vorobik, PhDSr. Policy Analyst . . . . . . . Emily

Roberson, PhDSo. California Regional Botanist . . .

Ileene AndersonInterim Rare Plant Botanist . . . Misa WardVegetation Ecologist . . Julie EvensVegetation Ecologist . . Anne KleinSan Bruno Mtn. Project Coordinator

Joe CannonLegislative Advocate .Vern GoehringLegal Advisor . . . . Sandy McCoyWebsite Coordinator . . . . . . . . . .

John DonaghueBulletin Editor . Michael Tomlinson

BOARD OF DIRECTORS

Carol Baird, Jim Bishop, VernGoerhing, Steve Hartman, DianaHickson, Lynn Houser, Lynne Kada,David L. Magney, Sandy McCoy, J.Spence McIntyre, Carol Witham(President)

CALIFORNIA NATIVE PLANT SOCIETYCALIFORNIA NATIVE PLANT SOCIETYCALIFORNIA NATIVE PLANT SOCIETYCALIFORNIA NATIVE PLANT SOCIETYCALIFORNIA NATIVE PLANT SOCIETY

MEMBERSHIPMembership form located on inside back cover; dues include

subscriptions to Fremontia and the Bulletin

Mariposa Lily . . . . . . . . . . . . $1,000Benefactor . . . . . . . . . . . . . . . . . $500Patron . . . . . . . . . . . . . . . . . . . . $250Plant Lover . . . . . . . . . . . . . . . . $100

Supporting . . . . . . . . . . . . . . . . . $75Family, Group, International . . . $45Individual or Library . . . . . . . . . $35Student/Retired/Limited Income . $20

CHAPTER COUNCILAlta Peak (Tulare) . . . . Joan StewartBristlecone (Inyo-Mono) . . . . . . . . .

Sherryl TaylorChannel Islands . . . . . . Lynne KadaDorothy King Young (Mendocino/

Sonoma Coast) . . . . Jon ThompsonEast Bay . . . . . . . . . Tony MoroscoEl Dorado . . . . . . . . . Amy HoffmanKern County . . . . . Laura StocktonLos Angeles/Santa Monica Mtns . . .

Betsey LandisMarin County . . . . . . . . Bob SoostMilo Baker (Sonoma County) . . . . .

Reny ParkerMojave Desert . . . . . . Tim ThomasMonterey Bay . . . . . . . . . . . . . . . . . .

Julie Anne DelgadoMount Lassen . . . . . . . . Jim BishopNapa Valley . . . . . . Marcie DannerNorth Coast . . . . . . . Larry LevineNorth San Joaquin . . . . Gail ClarkOrange County . . . . . Sarah JayneRedbud (Grass Valley/Auburn) . . . .

Richard HanesRiverside/San Bernardino counties . .

Katie BarrowsSacramento Valley . . Diana HicksonSan Diego . . . . . . . . Dave FlietnerSan Gabriel Mtns . . . . Lyn McAfeeSan Luis Obispo . . . . Dirk WaltersSanhedrin (Ukiah) . Chuck WilliamsSanta Clara Valley . . Georgia StigallSanta Cruz County . Janell HillmanSequoia (Fresno) . . . . Warren ShawShasta . . . . . . . . . . . Dave DuBoseSierra Foothills (Tuolumne, Cala-veras, Mariposa) . . . . Patrick StoneSouth Coast (Palos Verdes) . . . . . .

Barbara SattlerTahoe . . . . . . . . . . Michael HoganWillis L. Jepson (Solano) . . . . . . .

Allison FleckYerba Buena (San Francisco) . . . . .

Mark Heath

MATERIALS FORPUBLICATION

CNPS members and others are wel-come to contribute materials for pub-lication in Fremontia. See the insideback cover for manuscript submissioninstructions.

Prepress by ScanArt / Printed by Craftsman Trade Press

Linda Ann Vorobik, EditorKatheen Dickey, Convening Editor

Bob Hass, ProofreaderSusan Cotterel, Designer

Distributed in June 2004

CNPS, 2707 K Street, Suite 1; Sacramento, CA 95816-5113(916) 447-CNPS (2677) Fax: (916) 447-2727

[email protected]

PROGRAM DIRECTORS

CNPS Press . . . . . . . Holly Forbesand Gail Milliken

Conservation. . . . . David ChippingHorticulture. . . Peigi DuvallPosters . . . . . . Bertha McKinley and Wilma FolletteRare Plants. . . . . . Ann HowaldVegetation . . . . Todd Keeler-Wolf

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

CONTENTS

GUEST EDITORIALGUEST EDITORIALGUEST EDITORIALGUEST EDITORIALGUEST EDITORIAL ................................................................................................................................................................................................................................................................................................................................................................................................................................................................................. 22222

SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES SEAWEEDS TELL THEIR STORIES by Paul C. Silva ........................ 33333For every seaweed there are stories, such as those for two Pacific coast kelps, the sea palm(Postelsia palmaeformis) and the ephemeral laminaria (Laminaria ephemera). Paul Silvarelates the tales of their discovery and naming by Russian and Swedish explorers, keen-eyedVictorians, and resourceful Midwesterners.

CALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDSCALIFORNIA’S NON–NATIVE SEAWEEDS

by Kathy Ann Miller .................................................................................... 1010101010

It is well known that our terrestrial flora has numerous introduced species, some of which areespecially noxious. Less well known are the weeds of the seashore flora. Several of these are discussed,in particular Caulerpa taxifolia, which can be extremely damaging to seaweed habitats.

CALIFORNIA SEAWEED INDUSTRYCALIFORNIA SEAWEED INDUSTRYCALIFORNIA SEAWEED INDUSTRYCALIFORNIA SEAWEED INDUSTRYCALIFORNIA SEAWEED INDUSTRY by Paul C. Silva ................... 1616161616

Many of us are familiar with the red alga “nori,” which wraps around the rice in sushi. Fewof us, however, realize that seaweed products are used extensively in numerous industries,including the preparation of foods and beverages, the manufacture of cosmetics, pharmaceuti-cals, and ink, and in textile sizing.

DISTRIBUTION OF CALIFORNIA SEAWEEDSDISTRIBUTION OF CALIFORNIA SEAWEEDSDISTRIBUTION OF CALIFORNIA SEAWEEDSDISTRIBUTION OF CALIFORNIA SEAWEEDSDISTRIBUTION OF CALIFORNIA SEAWEEDS

by Paul C. Silva .............................................................................................. 1818181818

Seaweeds are restricted vertically by the availability of light in deep water and by their abilityto withstand desiccation in the intertidal zone. Each species has an optimal temperature rangewhich determines its geographic distribution. The role of Point Conception in separating cold-temperate and warm-temperate floras is explained in detail.

INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDSINTERTIDAL MONITORING OF CALIFORNIA SEAWEEDSINTERTIDAL MONITORING OF CALIFORNIA SEAWEEDSINTERTIDAL MONITORING OF CALIFORNIA SEAWEEDSINTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS

by Kathleen Dickey ....................................................................................... 2727272727

Marine biologists have documented the California intertidal for more than a century, butthere remains much to decipher. Threats to the intertidal, both natural and man-made, haveintensified efforts to monitor and further understand this fascinating ecosystem.

SEAWEED RESOURCESSEAWEED RESOURCESSEAWEED RESOURCESSEAWEED RESOURCESSEAWEED RESOURCES ......................................................................................................................................................................................................................................................................................................................................................................................................................................... 3030303030

HARVESTING AND COLLECTING SEAWEEDSHARVESTING AND COLLECTING SEAWEEDSHARVESTING AND COLLECTING SEAWEEDSHARVESTING AND COLLECTING SEAWEEDSHARVESTING AND COLLECTING SEAWEEDS ................................................................................................................................................................................................................................. 3131313131

NOTES AND COMMENTSNOTES AND COMMENTSNOTES AND COMMENTSNOTES AND COMMENTSNOTES AND COMMENTS ................................................................................................................................................................................................................................................................................................................................................................................................................ 3131313131

LETTERS TO THE EDITORLETTERS TO THE EDITORLETTERS TO THE EDITORLETTERS TO THE EDITORLETTERS TO THE EDITOR ........................................................................................................................................................................................................................................................................................................................................................................................................... 3232323232

BOOK RBOOK RBOOK RBOOK RBOOK REEEEEVVVVVIIIIIEEEEEWWWWW ............................................................................................................................................................................................................................................................................................................................................................................................................................................................................................................... 3232323232

F R E M O N T I A 1

THE COVER: Fronds and floats of Macrocystis pyrifera, the giant kelp, sometimes referred to as the “sequoia of the sea.”An individual kelp plant can grow up to two feet per day. Kelp forests provide habitat and food for a wide range of lifein central and southern California nearshore communities. Photograph courtesy of J. and J. Mondragon(www.mondragonphotos.com).

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

GUEST EDITORIAL

ur world is both challenging and inspiring, with moments of urgency juxtaposed with those of riveting beauty. The world of seaweeds is noexception. Last fall there was news of yet another old oil tanker off the coast ofSpain that was in danger of breaking and spilling its cargo. To add insult toinjury, this occurred along the same stretch of coast that had been inundatedwith oil just the year before. A request went out around the world via the AlgaeListserv (www.seaweed.ie) for baseline data and for assistance with tackling theformidable task of remediation. Despite continuous wake-up calls, we still seemto be “asleep at the wheel” with respect to protecting the intertidal regions ofthe world and preventing these disasters from recurring.

Fortunately, the scientific community has been “waking up” for nearly halfa century and is beginning to compile and synthesize long-term data sets thatwill document changes due to such events, and will thus provide ammunition inthe fight for laws and policies that will protect our coasts and their nativeecosystems, including the seaweed flora.

Worldwide this flora is impressive: there are some 8,000 species of marinealgae—simple, salt-water dwelling organisms that, to quote Dr. Mike Guiry,professor of phycology, “fall into the rather outdated general category of ‘plants.’ ”The algae include three major groups which have evolved quite separately and aredescribed by their accessory pigments: the reds (the most abundant), the browns(which include the kelps), and the greens (which may actually be part of the PlantKingdom).

Along the coast of California we have a particularly rich flora of seaweeds,which includes some 700 species. This great diversity is due in part to the cold,upwelling ocean waters that bring nutrients and oxygen to our coastal intertidalregions. In addition, the geological forces that have formed our coast resultedin a gorgeous patchwork of uplifted rock, providing a multitudinous array ofsubstrates for the seaweeds to anchor themselves with their holdfasts. Readabout the diversity of California seaweeds in Paul Silva’s article on biogeogra-phy on page 18. And, as in the terrestrial flora, some seaweeds found inCalifornia are natives of other regions. Kathy Ann Miller writes about weedyseaweeds in her article on page ten.

Seaweeds themselves boast some amazing records. Consider the following:They have been found as deep as 268 meters in the waters of the Caribbean, aswell as under the ice at Antarctica. The giant kelp, Macrocystis pyrifera, can growup to two feet per day. Recently a whole new group of red coralline seaweedshave been discovered called rhodoliths, individuals of which are at least 150years old, or which, if you take into account their capacity to reproducevegetatively, may be thousands of years old.

Whether we are aware of it or not, our lives are impacted daily by marinealgae, as they are used commercially in many familiar foods and products, suchas the thickening agent of toothpaste and paint, and as the emulsifier inchocolate milk. They are the basis of a thriving industry off the southernCalifornia coast. Paul Silva writes of seaweeds and industry in his article onpage 16.

Most native plant enthusiasts will celebrate with curiosity and appreciationCalifornia’s intertidal flora, but for those of you who might have less interest inthese salt-water dwellers, you are at first teased in with two stories of seaweeddiscovery (page three) by Paul Silva. Professor Silva deserves specialacknowledgement for his contribution to this issue, as he is one of the foremostauthorities on California seaweeds.

—Kathleen DickeyCNPS Santa Clara Valley Chapter

Member of the Fremontia Editorial Advisory Board

USEFUL WEBSITESUSEFUL WEBSITESUSEFUL WEBSITESUSEFUL WEBSITESUSEFUL WEBSITES

AND CONTACTAND CONTACTAND CONTACTAND CONTACTAND CONTACT

INFORMATIONINFORMATIONINFORMATIONINFORMATIONINFORMATION

Seaweed websites:See “Seaweed Resources,” p. 30

California Native PlantSociety (CNPS):www.cnps.org, with links toconservation issues, chapters,publications, policy, etc.To sign up for “NPCC News,”e-mail news on native plantscience and conservation, senda request to [email protected].

For updates on conservationissues:Audubon Society www.audubon.orgCenter for Biological Diversitywww.sw-center.orgNatural Resources DefenseCouncilwww.nrdc.orgSierra Clubwww.sierraclub.orgWilderness Societywww.wilderness.org

For voting information:League of Women Voterswww.lwv.org, includes online voterguide with state-specific nonparti-san election and candidateinformation.

US Senatewww.senate.gov

US House of Representativeswww.house.gov

California State Senatewww.sen.ca.gov

California State Assemblywww.assembly.ca.gov

To write letters:President George W. BushThe White House1600 Pennsylvania Ave. NWWashington, DC 20500

Senator Barbara Boxeror Senator Dianne FeinsteinUS SenateWashington, DC 20510

Your CA RepresentativeUS House of RepresentativesWashington, DC 20515

O

2 F R E M O N T I A

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

SEAWEEDS TELL THEIR STORIES

by Paul C. Silva

For every seaweed there is a story to tell, and getting to know these stories greatly enhances our appreciation of these diverseand beautiful plants. Here are two such stories.

F R E M O N T I A 3

SEA PALMSEA PALMSEA PALMSEA PALMSEA PALM

The western edge of the Ameri-can continent was put on the map,literally and figuratively, when, onthe 15th of July, 1741, the St. Paul,commanded by Alexei Chirikov,reached the shore of an island inwhat today we know as theAlexander Archipelago in southeast-ern Alaska. Vitus Bering, headingthe Second Kamchatka Expeditionand commanding the St. Peter, madelandfall at Cape St. Elias at the tipof Kayak Island a few days later (onJuly 18th). The two ships had pro-ceeded independently after havinglost sight of one another in the fog.

The St. Paul returned toKamchatka on October 8th, withmost of the crew dead from scurvy.The Saint Peter was undergoingeven greater hardships, beingwrecked on the westernmost side ofthe Commander Islands close toKamchatka. The crew overwinteredon this island, which they namedBering Island in memory of theircaptain, who had died shortly afterthe shipwreck. In the spring, enoughboards and nails were salvaged fromthe wreck to construct a small boat,which enabled only a handful ofsurvivors to return to Kamchatkaon August 25, 1742.

Tales of the great wealth ofnatural resources of “BolshayaZemlya” (“Big Land”), as the Rus-sians initially called Alaska, quicklyresulted in a flood of Russian ad-venturers, mostly trappers and furtraders. Their brutal treatment ofthe Aleut people and the near-ex-tinction of the sea otter are well-known dark episodes of history. By

Postelsia palmaeformis, the sea palm, was named after the Estonian geologist and artistwho painted seaweeds from nature during the 1826-1829 round-the-world voyage onthe vessel Seniavin. Illustration from F.J. Ruprecht, 1852: Neue oder unvollständigbekannte Pflanzen aus dem nördlichen Theile des stillen Oceans [New or incompletelyknown plants from the northern parts of the Pacific Ocean], St. Petersburg.

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

book, Illustrationes Algarum, con-tains the original descriptions ofmany species that are common inCalifornia, although no collectionswere made south of southeasternAlaska. The specific epithet,palmaeformis, is self-explanatory, theresilient plants bending over in thesurf reminiscent of palm trees dur-ing a hurricane or typhoon.

It may or may not be surprisingthat none of the several botanistswho sampled the marine flora ofMonterey during the late 18th cen-tury and early 19th century collectedthis quintessentially unique alga.Perhaps the reason for this lacunais that the sea palm, hanging on fordear life on wave-swept rocks, sig-nals would-be collectors to be onguard lest they find themselves in asimilar life-threatening position.

Vosnesensky provided amplecollection data. The three speci-mens illustrated in Ruprecht’s platewere collected on an islet at theentrance to Bodega Bay, called bythe natives “Omújüpai.” They calledthe sea palm “Kakgunu-chale,”whose meaning I do not know, butcertainly it would have nothing todo with palm trees.

The sea palm belongs to theorder Laminariales of the classPhaeophyceae (brown algae). It isone of several very distinctive kindsof kelp, as members of this orderare called. It is essentially an an-nual, growing rapidly in spring,

Postelsia palmaeformis, the sea palm,growing with red algae and gooseneckbarnacles. Photograph by G. Hansen.

In 1852 Ruprecht described one species of the surfgrass genus Phyllospadix(Zosteraceae Family). This plant is one of the few angiosperms that has adapted tolife in the intertidal surf. Photograph of flowers and leaves by S. Sharnoff.

1792 there was sufficient Russianpresence in Alaska to warrant theestablishment of a permamentsettlement on Kodiak Island, fol-lowed in 1799 by the founding ofthe town of Novoarkhangelsk (NewArkhangel, now known as Sitka).

In order to supply food for theRussian colony, the government, in1812, secured an area on theSonoma County coast, which wasplanted with crops and protectedby a fortification. It was namedRossiya (later called Fort Ross byAmericans) while a nearby river wasnamed Slavianka (later called Rus-sian River by Americans). Trappersand fur traders were soon raidingthe coast as far south as San Fran-cisco. Russians competed withAmericans for the eggs of sea birds,another lucrative item of trade. By1840 nature’s bounties were seri-ously depleted in northern Califor-nia, and the Russian government,feeling the hot breath of the Britishto the north and Mexicans to thesouth, decided to abandon theproject. The prestigious Academyof Sciences in St. Petersburg, uponhearing this decision, arranged fora young zoologist and naturalist,Ilya Vosnesensky, to go to Rossiyaand survey the nearby animals andplants. The Sonoma coast has manyspectacular kelps, and these werecollected by Vosnesensky, who ig-nored the smaller seaweeds. Hismission completed, the Russian gov-ernment closed Rossiya in 1842,

selling the fort and its contents toJohn Sutter. Vosnesensky by thistime was collecting in Baja Califor-nia. His collections were sent to theAcademy of Sciences, located in St.Petersburg.

The seaweeds collected byVosnesensky were studied by FranzJosef Ruprecht, who was born inAustro-Hungary, studied medicinein Prague, and became the curatorof botany at the Academy of Sci-ences at St. Petersburg in 1839. In1852 Ruprecht published a paper inwhich he treated only five seaweedsand one seagrass from SonomaCounty, but which is justly famousnot only for the three new generathat he described, but also for theoversized and remarkably lifelikelithographs. The plate for Egregiamenziesii is 1.46 meters long, haseight folds, and is spliced only once.

One of Ruprecht’s new generawas the sea palm, which he namedPostelsia palmaeformis. The genericname honors Alexander PhilipouPostels, an Estonian-born geologistand artist who had painted seaweedsfrom nature while on board theSeniavin during the voyage aroundthe world commanded by FriedrichLütke (also transliterated as Litke)in the period from 1826 to 1829. In1840, Postels and Ruprecht togetherhad published a monumental “el-ephant” folio volume treating thecollections of seaweeds made dur-ing that expedition. This very rare

4 F R E M O N T I A

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

maturing in late summer, discharg-ing spores, and remaining until thewaves destroy it by relentless pound-ing or knock it off its perch. In theintense competition for space in theintertidal zone, Postelsia often findsitself perched on mussels or bar-nacles, which may be swept off therocks all together. Overwintering isaccommodated by the microscopicsexual plants that develop from thedischarged spores. The fertilizedegg begins development of the erectplant, but then waits until earlyspring, when rapid growth occurs.The spores often stick to the sub-strate close to the mother plant, sothat in early spring a grove of verysmall “palm trees” may be seen sur-rounding the remains of the previ-

ous generation.At the same time that Ruprecht

described Postelsia, a Swedisharound-the-world expedition onboard the frigate Eugenie stoppedin San Francisco Bay, where thenaturalist, Nils Johan Andersson,collected a specimen of sea palm(probably from Point Lobos orLands End on the Pacific side ofSan Francisco). This specimen wasreferred to a Swedish botanist, JohnErhart Areschoug, who specializedin the study of kelps. Unaware ofRuprecht’s work, Areschoug de-scribed the new species as Virginiapalma-maris. The generic name maybring to mind the State of Virginiaor perhaps the Virgin Mary, but infact it commemorates the leader of

the expedition, C.A. Virgin. Theepithet, palmaeformis, again is self-explanatory. Areschoug’s paper waspublished in 1853, only a year afterRuprecht published Postelsia.

THE EPHEMERALTHE EPHEMERALTHE EPHEMERALTHE EPHEMERALTHE EPHEMERAL

LAMINARIALAMINARIALAMINARIALAMINARIALAMINARIA

Our second story, involving an-other distinctive kelp, begins on theMonterey Peninsula, nearly a half-century later. Laminaria ephemera,as the specific epithet implies, isephemeral, appearing in March anddisappearing in August. It is at-tached to the substrate by means ofa holdfast, from which there arisesa stem ending in a simple blade.The blade produces an abundanceof spores, which settle on a sub-strate and germinate to produceminute sexual plants. After an egghas been fertilized by a motile malegamete, the resulting zygote re-mains dormant or undergoes onlythe first stages of development intoa macroscopic plant during winter.Growth of the new plants is veryrapid in early spring, followed bymaturation, spore discharge, anddisintegration. Laminaria ephemerais one of only a few species of thegenus that are annual. In most spe-cies the macroscopic plant is peren-nial, the stem continuing to increasein diameter while the blade is re-newed at the beginning of eachgrowing season after being tornaway by the waves.

Laminaria ephemera is not onlyephemeral, but also elusive, appear-ing in large numbers at only a fewwidely separated sites. It thus es-caped the notice of naturalists onearly expeditions.

It did not escape the sharp eyesof Mrs. J.M. Weeks, however, whofirst recognized its distinctness dur-ing one of her frequent forays toCarmel Bay. Mrs. Weeks was anenthusiastic and dedicated seaweedcollector, as were several other la-dies living in Pacific Grove during

The life cycle of the bull kelp (Nereocystis) is typical of all kelps, including Postelsiapalmaeformis and Laminaria ephemera. Illustration by L. Scharf, from Seaweeds of thePacific Coast by J. Mondragon & J. Mondragon (Sea Challengers 2003).

F R E M O N T I A 5

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

Pressed specimens of Laminariaepehemera collected by Mrs. J.M.Weeks, whose persistence and carefulcollecting habits succeeded in gettingProfessor W.A. Setchell’s attention.Image use courtesy of the UniversityHerbarium, UC Berkeley.

6 F R E M O N T I A

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

the 1890s. Some were devoted tomaking spectacular mounts ondeckle-edged cardboard to be soldin curio shops, perhaps to end upon someone’s mantle in San Jose.Mrs. Weeks had a keen scientificinterest and rejoiced when shelearned that the University of Cali-fornia had added a phycologist toits faculty. That man was WilliamA. Setchell, who was brought toBerkeley in 1895 to be the head of anewly organized botany depart-ment. Setchell brought with himextensive knowledge of the algae,having studied under the tutelageof two eminent cryptogamic bota-nists, Daniel C. Eaton of Yale Uni-versity for his master’s degree, andWilliam G. Farlow of Harvard Uni-versity for his doctoral degree. Withthis background, he satisfied bothfactions on the faculty—Yalees andHarvardmen.

Arriving in Berkeley two monthsafter classes had begun, Setchell wasimpatient to sample the local flora.During Christmas break, heboarded a coastal steamer boundfor San Pedro with a stop in bothdirections at Port Harford (nowcalled Port San Luis). He collectedseaweeds at the two sites. Shortlyafter the Christmas break, Setchellreceived an introductory letter fromMrs. Weeks (dated February),which he promptly answered, andthus began a lively correspondencethat lasted through 1905.

The plant that was to be namedLaminaria ephemera was first men-tioned by Mrs. Weeks in a letterfrom Pacific Grove dated April 22,1897, written upon her return froman excursion to Carmel Bay madein a phaeton pulled by “Old Lottie”:“By this morns Express sent a cancontaining a Laminaria of which Ionce found, some three or four yearsa single specimen / I had begun tothink it a myth, as I could find nomore of it / Yestermorning I cameupon a great amount of it but mycontaining vessels & little cart werealready full / I could not secure

enough for a set.” (The “set” refersto the Phycotheca Boreali-Ameri-cana, a set of dried specimens soldto subscribers.) The remainder ofthe letter deals with possible ar-rangements whereby Setchell couldeither go to see the Laminaria orhave it collected for him by a resi-dent of Carmel. A year later, in aletter from “Carmel City” datedApril 13, 1898, Mrs. Weeks wrote:“The Laminaria I sent you lastspring is quite abundant. Do youcare to see any more of it? . . . Theplants are older than they were lastyear at this time. The stripes seemto be only an occasional feature /The sporangia cover a large por-tion of the frond. I noticed todayon some I gathered to send awaythe disk was smooth & round abouta fourth of an inch in diameter . . . Iwish you might see them / Theyseem of the Neriocystus [i.e.,Nereocystis] type . . .”A few days later(April 24), following Setchell’s re-ply, Mrs. Weeks wrote: “In regardto the Laminaria I sent you: I neednot say it is a painful disappoint-ment to me not to be able to get atthe plant. The portions of the beachthat had enough of it accessable arebedded in with sand and all but thetips are buried. The sand shifts in aday. I can see it out in the deeperwater but as the surf is bad there Idon’t feel strong enough to venture. . . I have been to a considerableexpense in coming down here & amnot very comfortable in the work. . . But I was so interested in thatLaminaria that I came. Why do youcall it Andersonii [Laminariaandersonii, a name misapplied to thespecies correctly called Laminariasetchellii]? It has no resemblance tothat plant. Its holdfast is differentfrom that, or indeed any other Lami-naria I have known / Its stipe istender / Its blade similar to theNeriocystus in substance and color.”

Mrs. Weeks persisted, impa-tiently waiting for Setchell to studyand describe this interesting sea-weed. In a letter from Santa Cruz

dated June 1, 1899, she wrote: “TheLaminaria I sent is not in any par-ticular like the one we have been inthe habit of calling L. Andersonia /It is not the color, thickness, norperiod of growth, or plan of form-ing sporangia of that plant . . . Ithought when I commenced gath-ering it was young Neriocystus andwondered it grew so long withoutan air bladder & when I found afully developed plant with severaldark, even bands of different widths[sporangial patches] running downthe length of it, it dawned upon methat I had found a stranger to myprevious collection . . . I showedone of my mounts to Dr. Anderson[a physician in Santa Cruz who waskeenly interested in seaweeds] & heasked if I had painted the stripes /He had never seen it. I was verymuch in hopes you would get downin time to see & study it where itgrew, for I think it is short lived”.

Professor W.A. Setchell, photographedin his office, circa 1934. Photograph byW. Matthews, courtesy of the UniversityHerbarium, UC Berkeley.

F R E M O N T I A 7

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

Letter from Mrs. Weeks to Professor Setchell, demonstrating that although Mrs. Weeks was not a professional botanist, she wasquite learned about seaweeds, not to mention classification and nomenclature.

Text of the letter reads as follows.

Martinez, Nov. 18, 1898

Prof. W. A. Setchell

Dear Friend,

I write to enquire in what subdivision Gloeosiphonia verticillaris and Lomentaria ovalis are? Have you assigned and named the membrane[sic] provisionally called Kalymenia reticulata & [ditto] corrugata? Is the Halidrys osmundaceae [ditto] Cystoseira? The Fucus fromSanta Cruz & the flat thin bladed Pelvetia that I sent from there, were they examined and what are they? Did you find more than oneoosphore in the flat Pelvetia[?] There is a plant growing abundantly in Carmel Bay that resembles the Gelidiums, Mr. Nott noticed it whenhe was there the first time. I will send a piece. I do not know what it is. Dr. Anderson thought it might be a Gymnogongrus. It is notdichotomous. I found on looking over things I had pressed free the rest of the Nitophyllum I showed you I will send a piece. Did I leave theFauschia with broad frond? If I did please save it for me.

Your obliged Friend,

J.M. Weeks

8 F R E M O N T I A

LEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCELEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCELEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCELEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCELEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCE

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

Receipt of a reprint of the paperin which Laminaria ephemera waseventually published by Setchell wasacknowledged by Mrs. Weeks onMarch 26, 1901, but it apparentlywasn’t until some months later (let-ter of November 25, 1901) that shediscovered with chagrin that in thesame paper Setchell had given thename Weeksia to a new genus of redalgae. Previously, when Setchell hadsuggested describing a new specieswith an epithet commemorating itsdiscoverer, Mrs. Weeks protested:“I am quite in earnest in not wish-ing my name attached to any plant”(letter of September 22, 1896). Mrs.Weeks summed up the feelings ofall amateur collectors: “I shall re-joice when definition and classifica-tion are definitely & clearly settled”(letter of June 26, 1899). Inciden-tally, any shortcomings in Mrs.Weeks’s writing should not be as-cribed to lack of education: shegraduated from Knox College inGalesburg, Illinois, in 1854, an un-usual achievement for a woman ofthat period. Her own explanationstrikes close to the heart of all ma-rine phycologists: “You must ex-cuse this if you can’t read it readilyas five days of rising about 3 A.M.& an ill turn make writing jerkybusiness” (letter of June 20, 1897).

The story of Laminaria ephem-era does not end here, but shifts toBritish Columbia, where JosephineE. Tilden, a professor in the botanydepartment at the University ofMinnesota, had established a bio-logical station on the southwestcoast of Vancouver Island with thefinancial help of one of her devotedstudents, Carolyn Crosby, whosefamily owned Crosby Mills (laterGeneral Mills). The station was nearPort Renfrew, a port used for ship-ping logs from the adjacent forest.It was an idyllic site, with massivegranite boulders covered with wave-beaten seaweed, including the seapalm (Postelsia). This seaweed wasparticularly striking, so that whenTilden arranged for the publica-

tion of a yearbook, she named it“Postelsia.” One of her students,Robert F. Griggs, found an abun-dance of an annual species of Lami-naria growing near the station, andunaware that it had already beendescribed and named by Setchell,proceeded to describe it exhaustivelyin the 1906 edition of “Postelsia.”He named it Renfrewia parvula.Griggs was impressed by the dis-coid holdfast vis-a-vis the stoutbranched holdfast of perennial spe-cies of Laminaria and thus estab-lished a new genus to accommo-date the local species and a previ-ously described Japanese species.The holdfast of Laminaria ephem-era, however, is not strictly discoid,often having rudimentary branches(haptera). Renfrewia currently is notrecognized as a genus separate fromLaminaria.

Although Laminaria ephemera isdelicate and shy, seeking shelteredpockets in the lowest intertidal andupper subtidal zones, while Postelsiapalmaeformis is rugged and resilient,defying the strongest of waves, theyshare similar stories of duplicated“discovery.” Even those seaweedsthat are less distinctive have storiesto tell. Victorians, who were in-trigued, even obsessed by the sea-side, its salubrious climate, and theinspiring array of creatures, weremoved to write poems, of which thefollowing, of uncertain origin butfound in M.M. Howard, “Oceanflowers and their teachings” (1846),is most appropriate:

Oh! call us not weeds, but flowersof the sea.

For lovely, and gay, and bright-tinted are we!

Our blush is as deep as the roseof thy bowers—

Then call us not weeds, we areOcean’s gay flowers.

REFERENCESREFERENCESREFERENCESREFERENCESREFERENCES

Areschoug, J.E. 1853.Virginia, ettnytt algslägte. Öfversigt afFörhandlingar Kongl. SvenskaVetenskaps-Akademien 10:145-147.

Griggs, R.F. 1906. Renfrewia parvula,a new kelp from Vancouver Island.Postelsia 1906:245-274.

Howard, M.M. 1846. Ocean flowers andtheir teachings. Binns and Goodwin.Bath, England.

Mondragon J. and J. Mondragon.2003. Seaweeds of the Pacific Coast.Sea Challengers. Monterey, CA.

Ruprecht, F.J. 1852. Neue oderunvöllstandig bekannte Pflanzen ausdem nördlichen Theile des StillenOceans. Mémoires de l’AcadémieImpériale des Sciences de Saint-Petérsbourg, Sixième Série, Sci-ences Naturelles 7(Bot.):55-82.

Setchell, W.A. 1901. Notes on algae.I. Zoe 5:121-129.

Professor Josephine E. Tilden returningfrom the third Pan Pacific ScienceCongress held in Tokyo, 1926. Use ofphotograph courtesy of S. Ducker.

F R E M O N T I A 9

Paul C. Silva, University Herbarium,UC, 1001 Valley Life Sciences Bldg. No.2465, Berkeley, CA [email protected]

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 41 0 F R E M O N T I A

he seaweed flora of Califor- nia is a world-class treasure. The magnificent diversity andabundance of seaweed populationsreflect the dramatic sweep of ourstate’s rich coastal environments andhabitats—from the Pacific North-west to the subtropics, includingrocky shores and reefs, sandybeaches, and offshore islands. Ourscientific knowledge of Californiaseaweeds is young, beginning withthe 1791 Malaspina expedition of dis-covery, growing through the effortsof European and east coast bota-nists, and coming of age with thecollected works of W.A. Setchell andN. L. Gardner at the University ofCalifornia at Berkeley. And yet, tothis day, the delineation of specieswithin our most common generaposes daunting challenges to phy-cologists, who continue to pursuefield, culture, and molecular studiesto unravel relationships at every taxo-nomic level.

The current comprehensive

flora, Marine Algae of California(Abbott and Hollenberg 1976) lists669 species of red, brown, and greenseaweeds. The actual number is cer-tainly much greater with the addi-tion of recently described species,species yet to be described, and spe-cies that are currently arriving. Ofthose more recent introductions,some come to California naturally,through range extensions due to ElNiño events and long-term climatechange, but some arrive via humanswho unintentionally introduce non-indigenous species on hulls of shipsor, for example, by improperly dis-posing of aquarium plants.

Non-native or exotic species,especially those that spread likeweeds, are both familiar and news-worthy. (Weeds, strictly defined, areplants that live where they are notwanted and are ecologically or eco-nomically harmful.) Since the be-ginning of civilization, humans havemoved plants and animals to servethem, either intentionally to pro-

vide food, shelter, or companion-ship, or inadvertently, as hitchhik-ers. For hundreds of years, our ter-restrial California has been gilded—and altered forever—by the unin-tentional introduction of Europeangrasses; the list of introduced Medi-terranean, South African, and Asianweeds is enormously long. Pampasgrass! Yellow star thistle! Scotchbroom! Fennel!

In the last few decades, we havecome to realize that our estuariesand coastal oceans are also vulner-able to introduced species, includ-ing invasive species that can causeeconomic and ecological night-mares. There are an estimated 175non-native species of plants and ani-mals in San Francisco Bay alone,and the number continues to grow.Rapacious invertebrates and fishlead the list and grab the news—theAsian clam, Chinese mitten crab,green crab, northern pike, stripedbass (an intentional introduction)are a few. But have you ever heard

CALIFORNIA’S NON-NATIVE SEAWEEDS

by Kathy Ann Miller

Caulerpa taxifolia, a beautiful, aggressive invasive tropical greenalga, appeared in a harbor near San Diego in 2000. Imagecourtesy of the University Herbarium, UC Berkeley.

T

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4 F R E M O N T I A 1 1

of the infamous “Killer Weed,”Caulerpa taxifolia? There are inva-sive seaweeds here, too.

Determining the presence ofexotic seaweed species can be diffi-cult if we have not witnessed theintroduction. We consider creaturesto be native when they occupy thearea in which they evolved; usuallythere are natural barriers to the dis-persal of a species. Sometimes thesebarriers shift through natural habi-tat or climate changes, allowing aspecies to expand its geographicrange. Sometimes we are able toaccess new areas via new technolo-gies, such as scuba diving orsubmersibles, and revise our recordsfor the ranges of species. But todistinguish introduced species fromthose that are native, we must relyupon the history of collection datacoupled with our understanding ofspecies distributions and natural dis-persal of populations.

When specimens from our coastwere sent back to European spe-cialists, vague collection notes (“inmare australe”) and mistakes assimple as mislabeling were frequent,and have left a legacy of confusion.For example, it was recently dis-covered and confirmed with mo-lecular data that the type locality(original collection) of a very com-mon red seaweed known as Iridaeacordata was the tip of South America,not Vancouver Island, Canada, theprovenance written on the her-barium sheet. That species name istherefore not applicable to our westcoast species (now known asMazzaella splendens, based on a Cali-fornia type specimen described bySetchell and Gardner). Similarly,the southern California rockweedHesperophycus californicus was onceknown as H. harveyanus, based on aplant supposedly collected inMonterey in the 1830s—but prob-ably collected in France and notHesperophycus at all.

Furthermore, many of our com-mon seaweeds have extraordinarilybroad “cosmopolitan” geographic

distributions and it is hard to dis-cern their history. These species areknown as “cryptogenic” becausetheir original provenances are hid-den from us. The ships that broughtearly explorers to our coasts werewooden reefs supporting creaturesfrom every port visited overmultiyear voyages, and were prob-ably significant sources of introduc-tions, long before scientists estab-lished a baseline for what is indig-enous. For example, Asparagopsistaxiformis is a red alga, originallydescribed from an Egyptian speci-men, but considered a cosmopoli-tan member of subtropical andtropical communities worldwide. Itoccurs in southern California (SanDiego, and the southern ChannelIslands). Is it introduced? Has itextended its range from Baja Cali-fornia, Mexico? Or has it “always”been here? We must critically re-visit nomenclatural history becausethe basis of our knowledge of sea-weed distributions is cultural as well

For this article and those thatfollow, common names are

only provided if available. Unliketerrestrial plants, most seaweedsdo not have colloquial names. Forpurposes of precision, a scientificname may be followed by itsauthor(s). If a species has been re-tained in its original genus, theauthor is the person who origi-nally described the species, for ex-ample, Postelsia palmaeformisRuprecht. If a species has beentransferred to another genus, thename of the describer will appearin parentheses followed by thename of the person who made thetransfer. Thus, the citationCaulerpa taxifolia (Vahl) C. Agardhindicates that this species was origi-nally described by Vahl (in 1802 asFucus taxifolius) and transferred toCaulerpa by C. Agardh (in 1817).

Label for only California collectionof Caulerpa taxifolia at the UniversityHerbarium, UC Berkeley. Herbariumlabels rarely include the colloquialname of the plant collected, but alwaysinclude the scientific name with itsauthor.

as biological.Finally, as every amateur and

professional phycologist knows,many species look extremely simi-lar and are difficult to tell apart,constituting complexes of what maybe cryptic (hidden) or sibling(closely related) species. Learningmore about our seaweed flora—anddetecting interlopers—is becomingincreasingly important.

The Mediterranean Sea, with85 introduced seaweeds, includingnine that are considered invasive, isa hot spot for introductions. At theThau Lagoon, an aquaculture siteon the southern coast of France, 45species of seaweeds (23% of theflora) are exotics, probably intro-duced with the Japanese oysterCrassostraea gigas. In contrast, we’verecorded a modest 12 non-nativeseaweed species (or 2% of the sea-weed flora) from our Californiacoast. Most of these are diminutiveand easily overlooked, and severalhave been reported only from our

Seaweed scientific names, common names,Seaweed scientific names, common names,Seaweed scientific names, common names,Seaweed scientific names, common names,Seaweed scientific names, common names,

and authorsand authorsand authorsand authorsand authors


own invasion hot spot, San Fran-cisco Bay.

Good examples are Aglaotham-nion tenuissima (Bonnemaison)Feldmann-Mazoyer and Polysiphoniadenudata (Dillwyn) Greville exHarvey, both small red filamentsoriginating in England and detectedin San Francisco Bay. More re-cently, another small red filamen-tous species, Polysiphonia harveyi J.Bailey, once thought to be a nativeof the east coast of North America,has been traced through molecularmethods as originating in Japan andspreading, via several introduc-tions, to the Atlantic, including theBritish Isles, New Zealand, and US,including California, where it hasmasqueraded as Polysiphoniaacuminata Gardner and P. simplexHollenberg. A great example of asibling/cryptic species complex.

Another example of a siblingspecies complex that is yet to beteased apart is the green algal ge-nus Bryopsis. In addition to Bryopsiscorticulans Setchell, threebryopsidoid entities, with differ-

ent reproductive characteristics, arerecorded from San Francisco Bay.One is entirely asexual, a good can-didate for an introduced species(no need to find a mate). And bythe way: Bryopsis hypnoidesLamouroux, reported fromHumboldt Co. to San Pedro, wasoriginally described from theMediterranean. Like so many Cali-fornia cryptogenic species with far-flung distributions, we have towonder: native or introduced?

The red alga Caulacanthusustulatus (Mertens ex Turner)Kützing is a cosmopolitan, crypto-genic species, reported from Japan,Indonesia, South Africa, Spain, NewZealand—all places where plantshave been designated as types forspecies now considered synonymsof C. ustulatus. This small, turfy spe-cies also occurs in Australia, Africa,India, and Hawaii, as well as BritishColumbia, Washington, and Cali-fornia. (As a student in 1979, I re-member being shown a small popu-lation on Vancouver Island. Exotic!Exciting!) In southern California,

Caulacanthus has been observed atnumerous sites during the last de-cade or so, but was not recordedduring extensive Bureau of LandManagement surveys 30 years ago.Just last year, I spotted a large redbrown patch in the intertidal as Iwalked my sister’s dog in a park onSan Francisco Bay. This proved tobe the first sighting of this speciesat the Bay. Where did it come from?How does it disperse? Are thesenew introductions, or is this speciesextending its range due to climatechange?

Lomentaria hakodatensis Yendois an inconspicuous red alga thathas quietly spread from its nativeJapan to the Mediterranean, Phil-ippines, Hawaiian Islands, Austra-lia and the west coast from BritishColumbia, Washington, and Or-egon to California and Mexico. Weassume it is an exotic because itwasn’t reported in early surveys(e.g., E. Yale Dawson’s surveys ofthe Northern coast from CapeMendocino to Crescent City). Be-cause many of our California spe-

Illustrations from the DeCew GuideIllustrations from the DeCew GuideIllustrations from the DeCew GuideIllustrations from the DeCew GuideIllustrations from the DeCew Guide

Throughout this article and within other articles in this Fremontiaissue, illustrations appear from Tom DeCew’s Guide to the Seaweeds

of British Columbia, Washington, Oregon, and Northern California, a Webversion of a book on northeast Pacific seaweeds conceived by the lateTom DeCew, and brought to its present state of completion with thecollaboration of Paul Silva, Richard Moe, and Robert Rasmussen.DeCew planned the book as a detailed complement to Marine Algae ofCalifornia by I.A. Abbott and G.J. Hollenberg (Stanford UniversityPress, 1976).

For each of the species of macroalga (seaweed) that occurs relativelycommonly in British Columbia, Washington, Oregon, or northernCalifornia, Decew assembled information from specimens in WestCoast herbaria and from personal collections. Each page contains agraphic depiction of distribution and reproductive status, often alongwith a specially commissioned original line drawing. He also compiledall published references of scientific studies for each species. The Webversion, which follows the layout of the book, including all of the dataand most of the illustrations for the species of brown and green algae,can be found at http://ucjeps.berkeley.edu/guide/.

Bryopsis hypnoides, a green alga, was described originally from the Mediterranean.Line drawing from DeCew Guide, used with permission from P. Silva.


cies also occur in Japan, it is diffi-cult to determine without good his-torical collections if a given speciesoccurs naturally throughout theeastern and western north Pacificor if it is an introduction. Gelidiumvagum Okamura is another smallred alga, native to Japan, China,and Russia. It has a limited distri-bution in British Columbia and hasnow appeared in Tomales Bay, Cali-fornia. Since Tomales Bay is hometo oyster mariculture, this speciesmay have been introduced as ahitchhiker with oyster spat fromWashington.

As in the Thau Lagoon, oystershave been the vector for many anintroduced species, including Sar-gassum muticum (Yendo) Fensholt,a species that can be considered ourfirst truly invasive species due to itsrapid spread. Introduced to the westcoast from Japan before World WarII, it was detected in British Co-lumbia in 1944, Oregon in 1947,Crescent City, California in 1963,Santa Catalina Island in southernCalifornia in 1970, and San Fran-cisco Bay in 1973. While studieshave not confirmed that Sargassummuticum is ecologically harmful,(e.g., displacing native species), it iscertainly a conspicuous and abun-dant part of our intertidal and shal-low subtidal communities. In 1973,when it arrived in the Solent regionof southern England (possibly froma shipment of British Columbia oys-ters), efforts to eradicate it ended infailure, even after volunteers re-moved 475 tons over the course ofthree years. Such efforts here, if wechose to undertake them 30 yearsafter the introduction, would surelyfail as well.

Oyster mariculture may be im-plicated in the dispersal of the greenalga Codium fragile subsp.tomentosoides (van Goor) P.C. Silva,another weed adept at asexual re-production. Codium fragile com-prises several well-behaved subspe-cies native to temperate Pacificcoasts, but was lacking in the Atlan-

tic until a weedy strain (subsp.tomentosoides) was introduced to theNetherlands from Japan (its pre-sumed source) shortly before 1900.This weed has spread with increas-ing rapidity throughout the NorthAtlantic, including the Mediterra-nean and the North American coastfrom Nova Scotia to North Caro-lina. It is now at home in NewZealand, Australia, and California(San Francisco and Tomales bays).An herbivorous sea slug checks thegrowth of the weed in New Zealand.In England and Ireland, the weedhas been reported to displace a na-tive species, Codium tomentosum. InNew England it overgrows and dis-lodges shellfish (thus earning thename “oyster thief”) and gangs upwith an introduced bryozoan to dis-place native kelps. To date in Cali-fornia it is confined to two bays andis thus separated from our nativesubspecies, which is well repre-sented on the outer coast fromAlaska to Baja California, Mexico.

The next two non-native

seaweeds are infamous aggressors,though how they will play in Cali-fornia is yet to be seen. In 2000,both the green alga Caulerpa taxifolia(M. Vahl) C. Agardh, and the brownkelp Undaria pinnatifida (Harvey)Suringar showed up in southernCalifornia. News of the first hasswamped interest in the second,because Caulerpa had provenitself to be a devastating plague inthe Mediterranean. Distasteful toherbivores, it has covered acres ofseafloor, out-competing nativeseagrasses and smothering nativefauna.

The culprit is an escaped inva-sive aquarium strain, identified asoriginating in Australia and subse-quently hybridizing with otherstrains, creating a strain tolerant oftemperate waters. During eelgrasssurveys, Caulerpa was discovered ina shallow lagoon, Agua Hedionda,near San Diego and in HuntingtonHarbor, California. Because theplant fragments and regenerates, itcannot be manually removed.

The distinctive “Dead man’s fingers,” Codium fragile subsp. tomentosoides, has thesecond, well-earned, name of “oyster thief.” Our native subspecies lives on the outercoast, while the invasive C. fragile subsp. tomentosoides has been found only in SanFrancisco and Tomales bays. Line drawing from DeCew Guide, used with permissionfrom P. Silva.


real until all species of Caulerpa arebanned from the aquarium trade.

While news of Caulerpa reachedthe public in newspapers, television,and Internet reports, Undariapinnatifida, a two meter long kelpnative to Japan, Korea, and Chinasilently skipped its way up the Cali-fornia coast from harbor to harbor,from Los Angeles to Santa Barbara.In 2001, it leaped to Monterey Har-bor and a small cove on the leesideof Santa Catalina Island. Over thepast several decades, Undaria has in-vaded Europe, the Mediterranean,South America, and Australia andNew Zealand, through deliberateintroductions (it is a prized food)and probably by hitchhiking on thehulls of ships. Like many invaders, ithas a broad tolerance for tempera-ture and thrives in quiet water, espe-cially in the presence of the nitrogenenrichment typical of harbors. InCalifornia harbors, it is a short-livedannual, recruiting in the early spring,and with a smaller pulse of newsporophytes in the fall at some sites,and then disappearing for the win-ter. Santa Catalina Island, one of theCalifornia Channel Islands, is theonly site at which it co-occurs withnative seaweeds in a relatively natu-ral (non-harbor) setting. There itlives at record-setting depths (up to24 m) in the clear offshore waters. Inthree years of observation at Catalina,Undaria has spread within the site,moving into shallower depths (in-cluding the understory of a giantkelp bed), but it has not extended itsrange to other sites around the is-land or other islands.

Because it is an annual, Undariadoes not appear to compete withperennial native kelps, but it cer-tainly warrants status as a potentialpest, capable of usurping habitat formuch of the summer. The Califor-nia Department of Fish and Gameis making efforts to eradicate it fromSanta Barbara harbor, and volun-teer groups are chipping away atthe Catalina and Monterey popula-tions. However, many predict that

Protect Our Coast from AnProtect Our Coast from AnProtect Our Coast from AnProtect Our Coast from AnProtect Our Coast from An

Invasive KelpInvasive KelpInvasive KelpInvasive KelpInvasive Kelp

U ndaria pinnatifida originated in Asia and has spreadaround the world to Australia, New Zealand, South America,

Europe—and now California. Undaria is unlike any other Californiakelp. Its blade is delicate (thin), elaborately lobed along the margin,and has a prominent midrib. It can be three to five feet long, and if youlook closely, you’ll see tiny dots scattered over the surface, formed bytufts of microscopic hairs. The reproductive structure (sporophyll)develops below the blade, justabove the holdfast, and is uniquein that it is deeply frilled, like old-fashioned ribbon candy. The bladepersists from spring through sum-mer, and then erodes back to theholdfast and sporophyll.

If you find this species, take apicture of it underwater. If this isnot possible, collect a specimenfor a photograph but do not throwit back into the water! Rather, dis-pose of it on land, and contact Dr.Kathy Ann Miller, Wrigley Ma-rine Science Center, SantaCatalina Island, (310) 510-4012([email protected]).

There are several websites withuseful additional informationabout this seaweed (search theWeb for Undaria pinnatifida). Forexample, the site at biology.ucsc.edu/people/potts/lonhart/undaria.htmlincludes many images of this plant.

eradication can be declared.Ten species and four subspecies

of Caulerpa are sold in aquariumstores in southern California. Ofthese, 12 are capable of surviving inthe temperate zone, and are poten-tial escapees.Because of difficultiesidentifying the invasive strain, thechance of a new introduction is very

Undaria pinnatifida, native toJapan, Korea, and China, is knownas wakame in Japanese cooking.Photograph by K.A. Miller.

Caulerpa patches were covered withtarp and bleach was injected beneath.A couple of years of this treatment(and several million dollars) resultedin a tentative victory: Caulerpa hasnot been seen at these sites in morethan a year, and has not been re-ported from new sites. Quarterlymonitoring continues until complete

Holdfast of Undaria pinnatifida.

Blade of Undaria pinnatifida.


any effort to remove Undaria at thispoint is too little and too late. Thekelp life history promotes long-termsurvival of the species throughhighly variable conditions. Eachkelp plant is capable of dispersingmillions of spores and establishingpopulations of the microscopic, fila-mentous sexual phase. Undaria isprobably here to stay, although weshould make every effort to eradi-cate it. Whether it will continue itsmarch up the coast remains to beseen.

A final species deserves men-tion, because there is evidence thatit may be establishing itself in Cali-fornia after years of unsuccessfulintroductions. This is the brownalga Ascophyllum nodosum (Linnaeus)Le Jolis, the knotted wrack nativeto the Atlantic, used to steam clamsand lobsters or pack bait. It has beensighted floating in the Bay on manyoccasions over the years, surely asdiscarded packing material, but onlythis year has a sample been collected

that might indicate that it has settledin. This too was floating, like theecad (ecological form) of Ascophyllumcalled scorpioides; attached plants havenot yet been observed.

This example reminds us thatthe establishment of a non-nativespecies is not a simple slam dunk.Those species blessed with a pecu-liar suite of attributes (broad physi-ological tolerances, copious repro-ductive abilities, and facile dispersalmechanisms) still require a large doseof luck to establish on foreign shores.

To date Sargassum muticum,Caulerpa taxifolia, Undaria pinnatifida,and perhaps Caulacanthus ustulatuscan be considered invasive weeds aswell as non-native components ofthe California flora. What will thefuture hold? Introductions seem tobe increasing (Caulerpa and Undariaarrived simultaneously in the newmillennium), and the consequencesof this global homogenization of spe-cies are mixed. Some, or perhapsmost, introductions are benign orneutral, but others are disastrous,setting the scene for “invasionalmeltdowns,” facilitating other inva-sions and sometimes increasing theimpact of subsequent invaders. I’veoften thought that the urchin-domi-nated habitats that have replacedsome of our kelp forests in southernCalifornia are fine settings for theinvasion of seaweed species distaste-ful to urchins, who could really chal-lenge our native kelps to a run fortheir money. But luck has been withus, so far. It’s clearly more impor-tant than ever to continue to studyour shores and to train our studentsto recognize our native Californiaseaweeds—and the new arrivals.


Abbott, I.A. and J. Hollenberg. 1976.Marine Algae of California. StanfordUniversity Press. 827 pp.

Boudouresque, C. F. and M. Verlaque.2002. Biological pollution in theMediterranean Sea: Invasive versus

introduced macrophytes. MarinePollution Bulletin 44:32-38.

Carleton, J.T. 2001. Introduced speciesin US coastal waters: environmentalimpacts and management priorites.Pew Ocean Commission, Arling-ton, Virginia. www.pewoceans.org

DeCew, T. (see sidebar on page 12)Guide to the Seaweeds of BritishColumbia, Washington, Oregon,and Northern California. Postedto the UC/JEPS website atucjeps.berkeley.edu/guide/

Deysher, L. and T. A. Norton. 1982.Dispersal and colonization in Sar-gassum muticum (Yendo) Fensholt.Journal of Experimental Marine Bi-ology and Ecology 56:179-195.

McIvor, Lynne, Christine A. Maggs,Jim Provan and Michael J.Stanhope. 2001. rbcL sequences re-veal multiple cryptic introductionsof the Japanese red alga Polysiphoniaharveyi. Molecular Ecology 10:911-919.

Silva, P.C. 1979. The benthic algalflora of central San Francisco Bay.Pp. 287-345 in San Francisco Bay:The Urbanized Estuary, T. J.Conomos, editor. Pacific Division,American Association for theAdvancement of Science. San Fran-cisco, CA.

Silva, P. C. 1996. California seaweedscollected by the Malaspina Expe-dition, especially Pelvetia (Fucales,Phaeophyceae). Madroño 43:345-354.

Silva, P. C., R. A. Woodfield, A. N.Cohen, L. H. Harris, & J. H.R.Goddard. 2002. First report of theAsian kelp Undaria pinnatifida inthe northeastern Pacific Ocean.Biological Invasions 4:333-338.

Verlaque, M. 2001. Checklist of themacroalgae of Thau Lagoon(Hérault, France), a hot spot ofmarine species introduction inEurope. Oceanologica Acta. 24:29-49.

Williams, S. L., and E. D. Grosholz.2002. Preliminary reports from theCaulerpa taxifolia invasion in south-ern California. Marine EcologyProgress Series 233:307-310.

The brown alga Ascophyllum nodosum,native to the Atlantic, is used to pack bait.It may be establishing itself in Californiaafter many years of unsuccessfulintroductions. Image courtesy ofUniversity Herbarium, UC Berkeley.

Kathy Ann Miller, Wrigley Marine Sci-ence Center on Santa Catalina Island,University of Southern California. 1 BigFisherman Cove, Avalon, CA [email protected]


Seaweed industry in Califor- nia was born of necessity. Prior to the First World

War, the mines of Stassfurt, Ger-many, were essentially the world’ssole source of potash, which is usedin making fertilizers, fireworks, andexplosives. To replace this sourcein wartime, the Hercules PowderCompany established a plant on theshore of San Diego Bay to obtainpotash (and other chemicals) by in-cinerating giant kelp (Macrocystissp.). At the peak of activity, therewere 1,100 employees, but compe-

tition from other sources of potashcaused the plant to close soon afterthe war ended.

The Second World War alsoproduced a crisis by cutting off oursupply of agar, normally importedfrom Japan. Agar is one of threecommercially important water-soluble polysaccharides found inseaweed. Although it is principallyknown as the gelling agent of thesubstrate on which bacteria aregrown, it also has many industrialuses. Prior to 1940, it was extractedmainly from various species of

CALIFORNIA SEAWEED INDUSTRY

by Paul C. Silva

Gelidium, a red alga. The first agarfactory in California was establishedin 1920 in Glendale, but operationswere soon moved to San Diego.Initially, the chief function of theindustry was to purify Japanese agarto bring it up to bacteriological stan-dards, but with the supply cut off bythe war, primary production wasessential. The source was Gelidiumrobustum, which could be obtainedin large quantities from fishermenin northern Baja California, whereit grows abundantly in the lowestintertidal and shallow subtidalzones. Agar production in SanDiego ceased about 1986 with theclosing of the American Agar andChemical Company, but the indus-try continues in Ensenada, BajaCalifornia.

The other two water-solublepolysaccharides (“phycocolloids”)are alginate (the sodium salt of alg-inic acid), obtained from large brownseaweeds (kelps and rockweeds), andcarrageenan, obtained from certainred seaweeds. All three phycocolloidsare emulsifiers and are often usedinterchangeably, but they differ incertain physical properties so thateach has its advantages. Both carra-geenan and alginate, more than agar,are used in the making of dairy prod-ucts, cosmetics, and pharmaceuticals,and in such industrial processes astextile sizing and ink manufacture.An alginate industry was started inSan Diego in 1927 and thrives todayunder the name ISP (InternationalSpecialty Products) San Diego. Theformer Kelco Company figuresprominently in the local history ofthis industry. The beds of Macrocystisthat extend from San Diego toMonterey provide the raw material.

A small factory for processingcarrageenan was operating in theSan Francisco Bay area during the

This Kelco ship harvests and processes the tops of the giant kelp, Macrocystispyrifera. The colloidal extract, sodium alginate, is used as a thickener in paints,dyes, beer, and canned foods. Use of photograph courtesy of J. Hansen.


1930s, using Mazzaella spp. as asource, but the world market is nowabundantly supplied, often satu-rated, by the output of seaweedfarms in many tropical countries,especially the Philippines, wherecarrageenan-rich Eucheuma spp. isgrown in ponds or on reefs close toshore. Historically, carrageenan wasextracted from Chondrus crispus(“Irish Moss”) by coastal peoples inthe North Atlantic and used tothicken puddings.

In Japan, commercial produc-tion of foodstuffs from seaweed, es-pecially amakusa nori fromPorphyra, kombu from Laminaria,and wakame from Undaria, is bigbusiness, but no such industry hasbeen established in California.

Along with commercial, large scale harvesting, Porphyra sp., known as nori, is collected in northern California as part of a cottageindustry.

A less utilitarian use of seaweed: toys made from bull kelp (Nereocystis luetkeana), withthe air bladders forming perfect doll heads. Use of photograph courtesy of K. Hubbard.



Benthic marine algae, or“seaweeds” as those of uswho know them on a first-

name basis prefer to call them, arethose algae that are attached torocks, other algae, animals, or man-made structures such as wood pil-ings. They are big enough to seeand feel, although it is necessary touse a hand lens or microscope todetermine the essential structure ofmany small species. While their eco-nomic importance is insignificantcompared to the role played by themicroscopic phytoplankton at thebase of the great ocean food web,their beauty and diversity havearoused the passion of naturalistsfor the past two centuries.

Seaweeds cling to the edge of

the continent, for the most part hid-den from ordinary view. Few peoplehave experienced the glorious sightof a rocky shore covered with a luxu-riant growth of seaweeds exposedat extreme low tide. In California,such a sight is possible only veryearly on summer mornings or justbefore dark in winter. These timesare dictated by the annual tidal cycleand are not likely to be chosen bythe casual visitor to the shore. Inaddition, most people head forsandy beaches, where they encoun-ter rotting masses of seaweedwashed ashore, often giant kelp(Macrocystis) intertwined withfeather boa kelp (Egregia), teemingwith hungry critters, such as kelpflies and sand hoppers. Children like

to pop the floats of Macrocystis anduse the Egregia as a jump rope. Thusthe seaweed experience of mostCalifornians would barely hint atthe glorious diversity and beauty tobe found in this fascinating groupof plants.

VERTICALVERTICALVERTICALVERTICALVERTICAL

DISTRIBUTIONDISTRIBUTIONDISTRIBUTIONDISTRIBUTIONDISTRIBUTION

Seaweeds do not venture up-ward beyond the splash zone be-cause they must be washed by thesea at least once each day. Few spe-cies can tolerate the long periodsof exposure imposed at high tidelevels. As the tide level decreases,the number of species increases

DISTRIBUTION OF CALIFORNIA SEAWEEDS

by Paul C. Silva

Corallina vancouveriensis, a coralline alga, is a common inhabitant of the mid to low intertidal. Coralline algae are impregnated withcalcium and may form a crust, be erect with jointed fronds, or both. Photograph by S. Sharnoff.


markedly, the greatest diversity be-ing found at the lowest tide level(about minus 0.6 meter on outercoasts of California) and in the ad-jacent subtidal fringe. Some specieslive only in the subtidal, at depthslimited by the availability of light.When the water is exceptionallyclear, a few species may grow atdepths as great as 268 meters, as inthe Bahamas (Littler et al. 1986). InCalifornia, the Channel Islands arenoted for the clarity of the sea, be-ing correlated with the absence ofriver runoff.

With increasing depth of water,the intensity of light decreasessharply while the quality of lightchanges dramatically. Red light iscompletely absorbed in the upperfew meters so that a blue-green twi-light prevails in deeper water. Thevarious classes of seaweeds (red,

brown, and green), each of whichhas a characteristic complement ofphotosynthetic pigments that is ef-fective in normal light, are adaptedto living in deep water by the pos-session of accessory pigments withappropriate absorption spectra.

At very low tide, the exposedshore often appears to be dividedinto several horizontal zones, dif-fering in color and texture depend-ing on the species that find optimalgrowth conditions at those particu-lar tide levels. Generally, one of thehigher zones is comprised of mem-bers of the Fucus family, which arecommonly called “rockweeds,”while the lowermost zone is domi-nated by members of the orderLaminariales, commonly called“kelps.” The subtidal fringe is oftenbright pink from encrusting calcar-eous algae of the order Corallinales.

FLORISTIC ANALYSISFLORISTIC ANALYSISFLORISTIC ANALYSISFLORISTIC ANALYSISFLORISTIC ANALYSIS

The coast of California is hometo about 670 species of marine al-gae, a mere 12% of the number ofhigher plant species. Endemics(those species restricted to Califor-nia) account for 14% compared to24% for higher plants (Hickman1993). By far the largest group ofspecies (44%) are those that are es-sentially restricted to the PacificCoast of North America, withranges extending from some pointnorth of California (some as farnorth as the Bering Sea) southward.This group includes mostly cold-temperate species, but also somethat survive or even thrive in mod-erately warm waters. Another group(21%) are those that are essentiallyrestricted to southern California andadjacent Baja California, constitut-ing a warm-temperate flora. Anequal number of species (21%) wereoriginally described from areasother than the North AmericanPacific Coast and include so-calledcosmopolitan species, introducedinvasive species (“weeds” in theusual sense of the word), and manyspecies with notably disjunct distri-butions that need to be confirmedby close taxonomic scrutiny. Manygenera and even some species ofthe California marine flora are be-lieved to occur also in Japan, butfuture molecular studies will prob-ably reveal at least slight geneticdifferences between western andeastern Pacific populations.

TEMPERATURE:TEMPERATURE:TEMPERATURE:TEMPERATURE:TEMPERATURE:

PRIMARY FACTOR INPRIMARY FACTOR INPRIMARY FACTOR INPRIMARY FACTOR INPRIMARY FACTOR IN

DETERMININGDETERMININGDETERMININGDETERMININGDETERMINING


Temperature is the overridingfactor in determining the geo-graphic distribution of seaweeds,while oceanic currents impose andmaintain the temperature regimes.The effect of the Gulf Stream on

Fucus gardneri, a rockweed, is a common inhabitant of the high intertidal zone. Linedrawing from DeCew Guide, used with permission from P. Silva.


climate is common knowledge. Itoriginates as the North EquatorialCurrent and passes through theStraits of Florida with sufficientvelocity to carry the warm wateralong the Atlantic coast of theUnited States as far as Cape Codand across the ocean to the BritishIsles and Norway. Equally spec-tacular, but in reverse, is the Peru-vian Current (formerly called theHumboldt Current), by which verycold water moving clockwise in theSouthern Ocean is deflected by thetip of South America, sending itnorthward along the Pacific coastand imposing temperate conditionsat tropical latitudes. Of greatestimportance in the North Pacific isthe Kuroshio Current system. Likethe Gulf Stream, it originates asthe North Equatorial Current andproceeds clockwise past the Phil-ippines and Taiwan along the coastof southern Japan, where it is calledthe Kuroshio Current. North ofJapan, it merges with the easterlydrift of the North Pacific Ocean.On reaching North America, partof the current is deflected north-ward in a counterclockwise gyre inthe Gulf of Alaska (Alaska Cur-rent) while part is deflected south-ward as the California Current.

The temperature of the cur-rents is sharply reduced under con-ditions favorable to upwelling. Inthis process, prevailing westerlywinds with cliffs to the left (in thenorthern hemisphere) push water,which has been warmed by the sun,along the coast. This forwardmovement is subjected to the Co-riolis force (an effect of the rota-tion of the earth), which causes thewarm water to be deflected to theright, away from the coast. Coldwater then wells up from moder-ate depths. Low-lying coasts allowthe winds to spread out and dimin-ish in strength, while headlandsconfine the winds and thus inten-sify the effect of upwelling. Innorthern California and northernBaja California, intense upwellingbegins in March and continuesthrough July, with water tempera-tures in the spring lower than thoseof winter. When the prevailingnorth-northwest winds diminishand upwelling ceases in autumn, arelatively warm surface counter-current (the Davidson Current)flows northward along the coast,being especially noticeable fromNovember through January. Thiscounter-current is generated by theCalifornia Current and flows in all

seasons, but is pushed downwardby upwelled water and moves up-ward when upwelling ceases.

In the southern hemisphere, theCoriolis force deflects the move-ment of water to the left, awayfrom the coast as in the northernhemisphere since the cliffs thatconfine the winds are on the right.Upwelling along the coasts of Chileand Peru lowers the temperatureof the Peruvian Current while up-welling in Namibia and AtlanticSouth Africa lowers the tempera-ture of the Benguela Current. Onthe west coast of both SouthAmerica and Africa, kelps, whichare quintessentially cold-waterplants, grow within the tropics.

Whether a current is warm orcold is relative. While the AlaskaCurrent is sufficiently warm toameliorate the climate of southernAlaska, the California current, en-forced by upwelling, is cold in itsnorthern reaches. Inshore watertemperatures reach their maximumin late August and early Septem-ber, but only in southern Califor-nia do they reach 20ºC or slightlyhigher. In northern California theusual annual range is 9-16ºC.

Whereas the distribution ofthose species that grow in the in-

Oceanic currents in the Atlantic Ocean.Note particularly the Gulf Stream, whichbrings warm water from the equator tothe eastern coast of North America andthe Atlantic coast of Europe.

Oceanic currents in the Pacific Ocean. Note particularly the Kuroshio Current system,which brings warm water from the equator to the coast of Japan but ends as therelatively cold California Current.


tertidal and shallow subtidal zonesis controlled by the temperature ofthe surface water, those that live indeeper waters usually have a par-ticular depth range correlated withtemperature. For example, the kelpAgarum fimbriatum grows in theshallow subtidal zone in the North-west, but it occurs in Californiaonly in deep water (to about 115meters) in the Channel Islands,being inhibited by the warm sur-face water.

SECONDARYSECONDARYSECONDARYSECONDARYSECONDARY

FACTORS INFACTORS INFACTORS INFACTORS INFACTORS IN

DETERMININGDETERMININGDETERMININGDETERMININGDETERMINING


When a species is within itsacceptable temperature range, itsdistribution may be limited by lackof a suitable habitat. Factors influ-encing suitability include availabil-ity of a firm substrate, exposure towave action, exposure to shiftingsand and gravel, and salinity. Somespecies (called cumaphytes, or surfplants) are restricted to the mostheavily exposed sites. Others pre-fer semi-protected reefs covered bysand. Most species have a particu-lar niche, such as surf grass beds atzero tide-level, or the vertical sidesof boulders. Most species can sur-vive short periods of lowered sa-linity, but few can tolerate consis-tently brackish water. For example,many conspicuous species com-monly found on the outer coast ofcentral California are missing inSan Francisco Bay, and those thatare there penetrate to a distancefrom the Golden Gate in directproportion to their tolerance ofbrackish water (Silva 1979). Wher-ever a creek discharges fresh waterinto the intertidal, the dischargewill be accompanied by a brackish-tolerant flora composed of brightgreen strands (Enteromorpha) andblackish green carpets (blue-greenprokaryotes, considered by some

biologists to be algae, and by otherbiologists to be bacteria).

GEOGRAPHICGEOGRAPHICGEOGRAPHICGEOGRAPHICGEOGRAPHIC


The biogeography of the ma-rine algae of California is a topicthat has engaged me for more than50 years. At the time that I begangraduate work (1946), there was awidely held belief that Point Con-ception caused a major break inthe marine biota, but there was nopublished evidence concerningseaweeds. The idea, which is largelyzoological, is based on the lifelongwork of William Healey Dall(1845-1927), a malacologist (Dall1916). I doubt that during Dall’slifetime any significant collectingof molluscs or any other form ofmarine life was done in the vicinityof Point Conception. The coast-line from Gaviota (just west ofSanta Barbara) northward toCarmel Bay was nearly inaccessibleprior to 1940. Highway 1 was builtbetween Carmel Bay and San LuisObispo County in the late 1930s,but was closed during the war.When I became immersed in thestudy of marine algae, I soon foundthat the only significant collectionsfrom along this stretch of coastwere those made by (and for)Rhoda Ramona Reed, a master’sstudent of Professor WilliamAlbert Setchell at Berkeley. Reed’sfamily owned a ranch near LionRock on the northwest shoulder ofPoint Buchon in San Luis ObispoCounty. Collections from thereand from “Oilport” were reportedin a thesis that was submitted inMay 1917 but was never published.Reed gave the location of herfather’s ranch as “Pecho,” and in-deed there is a Pecho Creek andPecho Rock nearby. “Oilport” wasa facility built in 1907 east of PortSan Luis in San Luis Obispo Bayat a site now occupied by ShellBeach. Inexplicably, “Oilport” is

Oceanic currents along the west coast ofNorth America. Note that the CaliforniaCurrent flows southward past Pt.Conception into northern Mexico beforereversing directions and sending a branchnorthward into the Southern CaliforniaBight.

neither in Goode’s California PlaceNames nor in Jepson’s index of geo-graphical names. Reed extendedthe range of several species hith-erto known only as far south asMonterey, and this informationwould have been useful to futureworkers had it been published. Inthe absence of published informa-tion on this stretch of coast, gener-alizations on the role of Point Con-ception in biogeography were infact based on comparisons betweenthe biota of Santa Barbara and thatof the Monterey Peninsula.

During the period 1946-1952,at the suggestion of Gilbert Mor-gan Smith, my major professor atStanford University, I collected in-tensively along the California coast,as well as on the Channel Islandsof southern California and themainland and coastal islands ofnorthern Baja California. It turnedout that Point Conception doesindeed play an important role inthe geographic distribution of Pa-cific coast marine algae (and assur-edly marine animals as well), but


Humboldt Countyunspecified

Acrosiphonia spinescens (Kützing) KjellmanPorphyra papenfussii KrishnamurthyPorphyra pseudolanceolata Krishnamurthy

TrinidadPalmaria hecatensis Hawkes

Cape MendocinoSphacelaria racemosa Greville

Mendocino CountyPoint Cabrillo (near Caspar)

Coilodesme bulligera StrømfeltWeeksia coccinea (Harvey) Lindstrom

Sonoma CountyShell Beach

Porphyra torta KrishnamurthyBodega Head and Bay

Cumathamnion sympodiophyllum Wynne &Daniels

Odonthalia washingtoniensis KylinPorphyra sanjuanensis Krishnamurthy

Marin CountyTomales Bay

Chaetomorpha cannabina (Areschoug)Kjellman

Fucus spiralis LinnaeusDuxbury Reef

Laminaria groenlandica RosenvingeOdonthalia lyallii (Harvey) J. AgardhPorphyra fucicola KrishnamurthyUlvaria obscura var. blyttii (Areschoug) Bliding

San Francisco CountySan Francisco (City) and San Francisco Bay

Acrosiphonia saxatilis (Ruprecht) VinogradovaElachista fucicola (Velley) AreschougLola lubrica (Setchell & Gardner) A. Hamel

& G. HamelPercursaria percursa (C. Agardh)

RosenvingePorphyra kanakaensis MumfordRalfsia fungiformia (Gunnerus) Setchell &

GardnerUrospora doliifera (Setchell & Gardner) Doty

Farallon IslandsMazzaella parksii (Setchell & Gardner)

Hughey et al.Ulva conglobata Kjellman

Santa Cruz CountyGreyhound Rock

Plocamium oregonum Doty

Monterey CountyMonterey Peninsula

Arthrocardia silvae JohansenFauchea fryeana SetchellHollenbergia subulata (Harvey) WollastonMonostroma grevillei (Thuret) WittrockPorphyra abbottiae KrishnamurthyPorphyra miniata (C. Agardh) C. AgardhPorphyra schizophylla HollenbergPylaiella tenella Setchell & GardnerRosenvingiella constricta (Setchell & Gardner)

SilvaSerraticardia macmillanii (Yendo) SilvaStictyosiphon tortilis (Ruprecht) Reinke

Point LobosChiharaea bodegensis Johansen

Malpaso CreekPorphyra smithii Hollenberg & Abbott

Kasler PointLessoniopsis littoralis (Tilden) ReinkePorphyra pulchra Hollenberg

Jade CoveLaminaria ephemera Setchell

Little Sur RiverIsabbottia ovalifolia (Kylin) Balakrishnan

Point SurPleurophycus gardneri Setchell & Saunders

Big Sur RiverUlva stenophylla Setchell & Gardner

San Luis Obispo Countyunspecified

Acrosiphonia coalita (Ruprecht) Scagelet al.

Porphyra lanceolata (Setchell & Hus) G.M.Smith

San SimeonHedophyllum sessile Setchell

Table 1. Southern endTable 1. Southern endTable 1. Southern endTable 1. Southern endTable 1. Southern end–points of conspicuous coldpoints of conspicuous coldpoints of conspicuous coldpoints of conspicuous coldpoints of conspicuous cold–temperatetemperatetemperatetemperatetemperate

seaweeds restricted to the mainland of California (but includingseaweeds restricted to the mainland of California (but includingseaweeds restricted to the mainland of California (but includingseaweeds restricted to the mainland of California (but includingseaweeds restricted to the mainland of California (but including

the Farallon Islands). Counties arranged from north to south.the Farallon Islands). Counties arranged from north to south.the Farallon Islands). Counties arranged from north to south.the Farallon Islands). Counties arranged from north to south.the Farallon Islands). Counties arranged from north to south.


not in the simplistic way the storywas usually told.

At Point Conception, the coastveers sharply eastward towardSanta Barbara. The relatively coldCalifornia Current continues itssouthward sweep, bathing thenorthern Channel Islands and thenflowing into Mexican latitudeswhere it is warmed. Part of it thenreturns northward, bathing thesouthern Channel Islands with rela-tively warm water and continuingcounter-clockwise in the SouthernCalifornia Bight. The other partcontinues southward as a weak cur-rent. The relatively warm water inthe Bight creates a sharp tempera-ture gradient immediately east ofPoint Conception so that theSanta Barbara flora is indeed verydifferent from that of the MontereyPeninsula.

The ranges of cold-temperate

CayucosPorphyropsis coccinea (Areschoug) Rosenvinge

Atascadero State BeachBornetia californica Abbott

Montana de Oro State ParkHymenena multiloba (J. Agardh) KylinKornmannia leptoderma (Kjellman) BlidingMacrocystis integrifolia BoryPercursaria dawsonii Hollenberg & AbbottPylaiella gardneri Collins

Lion RockPelvetiopsis limitata (Setchell) GardnerPostelsia palmaeformis Ruprecht

Diablo CoveAlaria marginata Postels & RuprechtCallophyllis crenulata SetchellClathromorphum parcum (Setchell & Foslie)

AdeyNereocystis luetkeana (Mertens f.) Postels &

RuprechtShell Beach

Hymenena fryeana KylinPismo Beach

Farlowia conferta (Setchell) Abbott

Santa Barbara CountyPoint Sal

Dictyoneurum californicum RuprechtErythrophyllum delesserioides J. Agardh

Point ConceptionAhnfeltiopsis linearis (C. Agardh) Silva &

DeCewAnalipus japonicus (Harvey) WynneBossiella plumosa (Manza) SilvaConstantinea simplex SetchellDilsea californica (J. Agardh) KuntzeFucus gardneri SilvaHalosaccion glandiforme (S.G. Gmelin)

RuprechtHymenena flabelligera (J. Agardh) KylinMastocarpus jardinii (J. Agardh) J. WestOdonthalia floccosa (Esper) FalkenbergPhaeostrophion irregulare Setchell &

GardnerGaviota

Gloiosiphonia verticillaris FarlowGoleta

Palmaria mollis (Setchell & Gardner) vander Meer & Bird

Table 1.Table 1.Table 1.Table 1.Table 1. (cont.) (cont.) (cont.) (cont.) (cont.)

The kelp Dictyoneurum califor-nicum is common between CapeMendocino and Monterey, but isfound as far north as BritishColumbia and as far south as SantaBarbara County. Line drawingfrom DeCew Guide, used withpermission from P. Silva.


species are related to temperaturerather than to latitude. Many cold-temperate species drop out at vari-ous points along the coast even be-fore they reach Point Conception(see Table 1). Postelsia palmaeformis,for example, occurs only as far southas Lion Rock in San Luis ObispoCounty. Some species drop out atPoint Conception or intrude a shortdistance into the Bight. Many spe-cies, however, continue their distri-bution on the northern Channel Is-lands (see Table 2). Some of thesespecies skip the southern ChannelIslands but appear again on themainland of northern Baja Califor-nia in pockets of upwelled cold wa-ter (see Table 3). Laminaria setchellii,for example, occurs on the Califor-nia mainland as far south as PointConception, then on the northernChannel Islands, with the south-ernmost locality being Punta Banda,just south of Ensenada, Mexico.Some species skip the Channel Is-lands as well as southern Califor-nia, reappearing in upwelling areasof northern Baja California (seeTable 4).

The warm-temperate flora ofsouthern California does not offera mirror image of the pattern shownby the cold-temperate flora in whichspecies drop out progressively asthe water temperature increases.Rather, it is largely confined to wa-ters south of Point Conception, withonly a few species venturing farthernorth. Those that do venture north-ward are restricted to pockets ofrelatively warm water, such asStillwater Cove in Carmel and atSanta Cruz (see Table 5). At LaJolla and in the southern ChannelIslands (San Clemente and SantaCatalina islands) there are severalspecies that may be found in nearlyall subtropical and tropical waters.As pointed out by Abbott andHollenberg (1976), the marine floraof southern California comprisesmore species but less biomass thanthat of northern California, as isusually the case when a cold-water

including a remarkable number ofmembers of the order Rhody-meniales, are totally subtidal accord-ing to data given by Abbott andHollenberg (1976). Their verticalranges are variable. A few speciesoccur in the subtidal fringe just be-low the lowest tide level and extenddownward, but the majority are re-stricted to depths of at least tenmeters. A few species occur as deepas 100 meters. Until the advent of

flora is compared with a warm-water flora. Throughout the world,large fleshy brown and red algae aregenerally restricted to cold water.

An excellent overview of theseaweeds of southern California isgiven by Murray and Blair (1993),who provide floristic, geographic,and ecological analyses, including adetailed account of communitystructure and productivity. Nearly60 species of California seaweeds,

Table 2. Conspicuous coldTable 2. Conspicuous coldTable 2. Conspicuous coldTable 2. Conspicuous coldTable 2. Conspicuous cold-temperatetemperatetemperatetemperatetemperate

seaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland end-points betweenpoints betweenpoints betweenpoints betweenpoints between

the Monterey Peninsula and Goleta, Santathe Monterey Peninsula and Goleta, Santathe Monterey Peninsula and Goleta, Santathe Monterey Peninsula and Goleta, Santathe Monterey Peninsula and Goleta, Santa

Barbara County, but also found on theBarbara County, but also found on theBarbara County, but also found on theBarbara County, but also found on theBarbara County, but also found on the

Channel Islands.Channel Islands.Channel Islands.Channel Islands.Channel Islands.

Antithamnionella spirographidis (Schiffner) WollastonHalymenia schizymenioides Hollenberg & AbbottMazzaella flaccida (Setchell & Gardner) FredericqMazzaella splendens (Setchell & Gardner) FredericqNeoptilota hypnoides (Harvey) KylinNeorhodomela oregona (Doty) MasudaPorphyra variegata Setchell & HusPrasiola meridionalis Setchell & GardnerUrospora penicilliformis (Roth) Areschoug

Hedophyllum sessile, a distinctive kelp, is found from Alaska to Point Conception, buthas yet to be found between Santa Cruz and Point Sur. Line drawing from DeCewGuide, used with permission from P. Silva.


scuba diving, several species wereknown only from plants cast ashoreafter storms or pulled up in dredgehauls. A few species have been ob-served by divers, but are rarely ifever cast ashore, presumably disin-tegrating in situ. The geographicranges of these subtidal species areerratic, in part depending on theavailability of certain areas for sam-pling. Many ranges encompass oneor more of several areas favorableto scuba diving and dredging. Theseareas include Bowie Seamount(about 75 miles west of the QueenCharlotte Islands, British Colum-bia), the Puget Sound (particularlyHein Bank in the San Juan Islands),Cordell Bank (Marin County), theFarallon Islands (San FranciscoCounty), Monterey and Carmelsubmarine canyons and SchmiederBank (Monterey County), DiabloSubmarine Canyon (San LuisObispo County), Scripps and LaJolla submarine canyons (San Di-ego County), and various diving anddredging sites in the Channel Is-lands and northern Baja California.including Tanner and Cortes banks.Kelp canopies, particularly those ofMacrocystis pyrifera, provide a habi-tat for a specialized algal flora to-gether with associated fishes andinvertebrates.

A very few species of the Cali-fornia flora are insular endemics,occurring in the southern ChannelIslands as well as on various Mexi-can islands, especially IslaGuadalupe and Rocas Alijos. Thehigh degree of endemism of themarine biota of Isla Guadalupe iseasily explained by the ecologicalisolation of this island resulting fromits position to the west of the coldupwelled waters at the same lati-tude on the Baja California penin-sula. The contrast between the ma-rine biota of Isla Guadalupe andthe mainland, which is only about160 nautical miles to the east, isstriking. The absence of these insu-lar endemics on the southern Cali-fornia mainland is more difficult to

explain. Part of the explanation maybe the deterioration of mainlandhabitats. These endemics may haveat one time been found on the main-land. The occurrence of several spe-cies of seaweeds that were found atSan Pedro a hundred years ago, butnot in recent times, is documentedby specimens in the herbarium atthe University of California, Ber-keley. Alternatively, these endemicsmay be represented by small subtidalcoastal populations waiting to bediscovered. Three conspicuous spe-cies are currently considered strictlyinsular: Cystoseira neglecta Setchell& Gardner and Sargassum palmeriGrunow (both brown algae) and anundescribed species of Codium (agreen alga).

The Monterey Peninsula is re-markably rich in seaweeds, both innumber of species and in luxuri-ance of growth. Its richness in part


seaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland end-points inpoints inpoints inpoints inpoints in

California between the MontereyCalifornia between the MontereyCalifornia between the MontereyCalifornia between the MontereyCalifornia between the Monterey

Peninsula and Goleta, Santa BarbaraPeninsula and Goleta, Santa BarbaraPeninsula and Goleta, Santa BarbaraPeninsula and Goleta, Santa BarbaraPeninsula and Goleta, Santa Barbara

County, but also found on the ChannelCounty, but also found on the ChannelCounty, but also found on the ChannelCounty, but also found on the ChannelCounty, but also found on the Channel

Islands and in northern Baja California.Islands and in northern Baja California.Islands and in northern Baja California.Islands and in northern Baja California.Islands and in northern Baja California.

Bossiella californica (Decaisne) SilvaCalliarthron tuberculosum (Postels & Ruprecht) DawsonCallocolax fungiformis KylinCallophyllis obtusifolia J. AgardhChondracanthus corymbiferus (Kützing) GuiryDesmarestia latifrons K¸tzingFarlowia compressa J. AgardhGloiopeltis furcata (Postels & Ruprecht) J. AgardhHaplogloia andersonii (Farlow) LevringLaminaria setchellii SilvaMastocarpus papillatus (C. Agardh) J. WestNeorhodomela larix (Turner) MasudaPeyssonneliopsis epiphytica Setchell & LawsonSchimmelmannia plumosa (Setchell) AbbottScytosiphon dotyi WynneSoranthera ulvoidea Postels & Ruprecht

Hesperophycus californicus, a conspicuousbrown warm-water rockweed, is onlyrarely found north of Santa Cruz. Linedrawing from DeCew Guide, used withpermission from P. Silva.



seaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland endseaweeds with mainland end-points inpoints inpoints inpoints inpoints in

California between the Monterey PeninsulaCalifornia between the Monterey PeninsulaCalifornia between the Monterey PeninsulaCalifornia between the Monterey PeninsulaCalifornia between the Monterey Peninsula

and Gaviota, Santa Barbara County, andand Gaviota, Santa Barbara County, andand Gaviota, Santa Barbara County, andand Gaviota, Santa Barbara County, andand Gaviota, Santa Barbara County, and

also found in northern Baja Californiaalso found in northern Baja Californiaalso found in northern Baja Californiaalso found in northern Baja Californiaalso found in northern Baja California

but not in the Channel Islands.but not in the Channel Islands.but not in the Channel Islands.but not in the Channel Islands.but not in the Channel Islands.

Ahnfeltiopsis pacifica Silva & DeCew, ined.Blidingia minima (Naegeli) KylinDelesseria decipiens J. AgardhDesmarestia viridis (O.F. Mueller) LamourouxPorphyra gardneri (Smith & Hollenberg) HawkesPrionitis filiformis KylinPtilota filicina J. Agardh

Table 5. Northern endTable 5. Northern endTable 5. Northern endTable 5. Northern endTable 5. Northern end-points ofpoints ofpoints ofpoints ofpoints of

conspicuous warmconspicuous warmconspicuous warmconspicuous warmconspicuous warm-temperate seaweeds.temperate seaweeds.temperate seaweeds.temperate seaweeds.temperate seaweeds.

Santa CruzChondria decipiens KylinChondria nidifica HarveyHesperophycus californicus Silva

Monterey PeninsulaAcrosorium venulosum (Zanardini) KylinHelminthora stricta GardnerLaurencia pacifica KylinLaurencia subopposita (J. Agardh) Setchell

San Luis Obispo County Cambria

Leptocladia binghamiae J. Agardh Shell Beach

Cystoseira setchellii Gardner

Los Angeles CountyRedondo Beach

Laurencia snyderae Dawson

port of sand, and diversity of sub-strate. An exemplary account of thehabitats and distribution of theseaweeds and seagrasses of a par-ticular region is that by Stewart(1991), who shares in satisfyingdetail her extensive long-term ob-servations of the marine flora ofSan Diego County.


Abbott, I.A. and G.J. Hollenberg.1976. Marine Algae of California.Stanford University Press.Stanford, CA. 827 pp.

Dall, W.H. 1916. Checklist of the Re-cent Bivalve Mollusks (Pelecypoda) ofthe Northwest Coast of America Fromthe Polar Sea to San Diego, Califor-nia. Southwest Museum, Los An-geles, CA. 44 pp.

Hickman, J.C. (ed.) 1993. The JepsonManual: Higher Plants of California.University of California Press. Ber-keley, CA. 1400 pp.

Littler, M.M., D.S. Littler, S.M. Blair,and J.N. Norris. 1986. Deep-wa-ter plant communities from an un-charted seamount off San SalvadorIsland, Bahamas: Distribution,abundance, and primary productiv-ity. Deep-Sea Research 33:881-892.

Murray, S.N. and R.N. Bray. 1993.Benthic macrophytes. In Ecology ofthe Southern California Bight, ed.M.D. Dailey, D.J. Reish, and J.W.Anderson, pp. 304-368. Universityof California Press. Berkeley, CA.

Silva, P.C. 1979. The benthic algalflora of central San Francisco Bay.In San Francisco Bay: The UrbanizedEstuary, ed. T.J. Conomos, pp. 287-345. Pacific Division, American As-sociation for the Advancement ofScience, San Francisco, CA.

Stewart, J.G. 1991. Marine algae andseagrasses of San Diego County. Cali-fornia Sea Grant College ReportNo. T-CSGCP-020. 197 pp.

is due to the equally remarkablevariety of habitats in the area, sothat certain species find the lack ofa suitable habitat rather than highertemperatures a barrier to extend-ing their range southward (seeTable 1).

In any given area, the general-

ized factors that affect the distri-bution of seaweeds are alwaysmodified, to a significant degree incertain places and at certain times,by local conditions such as inshorecurrents and countercurrents, lo-calized upwelling, seasonal riverand creek discharge, seasonal trans-



embers of the CaliforniaNative Plant Society have long been passionate

pioneers in documenting the iden-tity, distribution, and abundance ofterrestrial plants. Botanists world-wide estimate that more than halfof all the plants in the world havenow been identified. Marine plantsand systems have also been wellstudied, and more than 8,000 spe-cies of marine macroalgae have beendocumented worldwide. In Califor-nia alone, more than 650 speciesof marine algae are known, butthere remains much to decipherconcerning patterns and changes incommunity structure and speciesrichness. More research is especially

needed to correlate changes in theseaweed flora and vegetation struc-ture with external forces, bothphysical and anthropogenic. Oneprime lesson gleaned from over halfa century of intertidal monitoringis that the inherent patchiness inintertidal community organizationmakes extrapolation from observedchanges to supposed causes chal-lenging, and thus there is a greatneed for careful documentation.

Intertidal monitoring studies fallinto three general categories: 1) thedescriptive or “snap-shot” studies;2) longer term studies, includingexperiments testing questions thatarose from the initial studies; and 3)long-term, collaborative monitor-ing projects dedicated to discerning

and describing patterns over time.These studies have been punctu-ated and inspired by external eventssuch as oil spills, nuclear power plantdevelopment, sewage outfalls, andother impacts of human populationgrowth.

It is challenging to census themarine environment consistentlyand over time, chiefly due to a lackof funding and personnel for such along-term commitment. It is alsooften a physically demanding, oreven hazardous job—there is occa-sionally the possibility of being bit-ten by an angry elephant seal in thecourse of a count! A benefit of thefocus on documention of intertidalhabitats and species has been therenewed appreciation of and need

INTERTIDAL MONITORING OF

CALIFORNIA SEAWEEDS

by Kathleen Dickey

Teaching census techniques is an important part of the continuity of long-term monitoring projects. Here a group of students fromthe Branson School in Marin County are monitoring Duxbury Reef through the Sustainable Seas program, working with the Gulfof the Farallones National Marine Sanctuary. Photograph courtesy of J. Saltzman, Farallones Marine Sanctuary Association.

M

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

for well-trained taxonomists, natu-ralists, and organismal biologists.After all, it’s no use counting some-thing that you can’t identify.

A BRIEF HISTORYA BRIEF HISTORYA BRIEF HISTORYA BRIEF HISTORYA BRIEF HISTORY

The first surveys done in theintertidal zone were primarily de-scriptive. Beginning in the 1930sthe early studies (notably those doneby W.G. Hewatt, E. Yale Dawson,P.K. Dayton, M.S. Foster, M.M.Littler, and S. Murray—for a re-view of early work see Murray 2001)were carefully executed, using clearand readily repeated methodology.The locations of many of the earlystudies were determined by accessto the tidepools and proximity toresearch stations.

E.F. Ricketts, author of theseminal Between Pacific Tides andproprietor of a biological supplyhouse (not to mention the inspira-tion for the main character in JohnSteinbeck’s novel Cannery Row),

intensively documented the rockyintertidal inhabitants and their re-spective zones on the MontereyPeninsula. One of the earliest for-mal intertidal censuses was con-ducted in a benchmark study in thesame area by W.G. Hewatt whenhe set up a permanent transect tosurvey intertidal invertebrates atHopkins Marine Station in 1931-33. Unfortunately for the seaweedcommunity, he didn’t includeplants. Additional survey work wasdone by E. Yale Dawson, who moni-tored 42 transects in southern Cali-fornia in 1963.

In 1973, John Pearse and hisstudents, funded by Sea Grant andlocal government agencies, con-ducted a survey of a mudstone reefusing meter-square plots along per-manent vertical transects (goingfrom the high tide to low tide zone).They established one transectbeneath a sewage outfall that wasclosed down in 1976. They havecontinued to monitor these sites.

WAKE–UP CALLSWAKE–UP CALLSWAKE–UP CALLSWAKE–UP CALLSWAKE–UP CALLS

AND THEIRAND THEIRAND THEIRAND THEIRAND THEIR

CONSEQUENCESCONSEQUENCESCONSEQUENCESCONSEQUENCESCONSEQUENCES

In 1969 an oil platform blewout in the Santa Barbara Channel.The ensuing disaster prompted aseries of studies, sponsored by theMinerals Management Services, inorder to obtain baseline data on thesouthern California coast. Suddenlythe surveys that had occurred in theprevious four decades had greatlyincreased value.

In contrast to the Santa Barbaraoil spill, which stirred both localand statewide interest, the 1989Exxon Valdez disaster grabbed at-tention around the world, whensome eleven million gallons of oilspilled into Prince William Sound,Alaska, fouling 1,300 miles of a here-tofore pristine intertidal region.When it came time for Exxon tofulfill their legal obligation toremediate these Alaskan habitatsafter the disaster, quantitative datawere insufficient to establish what

2 8 F R E M O N T I A

Recognizing the differences in branching patterns between two species of Microcladia as well as the differences in habit is criticalto an accurate census. Microcladia borealis (left, x2) is saxicolous (grows on rocks) whereas M. coulteri (right, x 1/2) is epiphytic (growson other plants). Line drawings from the DeCew Guide, used with permission from P. Silva.


had been lost. Unlike California,where there had been previous ef-forts along the southern and cen-tral coast to collect intertidal data,there simply wasn’t enough knownabout the species and communitiesin areas affected by the Alaskan spill.As a consequence, Exxon is nowappealing the settlement for puni-tive damages. This case providedan urgent wake-up call to those whomonitor the marine environment.If the acquisition of basic scientificknowledge was not a strong enoughincentive for establishing baselinedata, then surely economics mightprovide a convincing argument.

WHAT HAVE WEWHAT HAVE WEWHAT HAVE WEWHAT HAVE WEWHAT HAVE WE

LEARNED?LEARNED?LEARNED?LEARNED?LEARNED?

In some instances, scientistshave documented considerablechange. In other cases the resultshave been ambiguous. In follow-upsurveys in the rocky intertidal ofsouthern California localities, SteveMurray and his students have ob-served that the relative abundanceof highly productive foliose algaehas declined, to be replaced by turf-forming seaweeds, corallines, andcrusts. The reasons for this may beanthropogenic, as there are simplymany more people coming to thesouthern shores than before.In fact, it is estimated that over thepast three decades the population ofcoastal counties in southernCalifornia has increased by morethan 50%.

In 1995, 60 years after theHewatt census of invertebrates atHopkins Marine Station, Barry etal. repeated the same work, usingthe same transect and plots. Theywere surprised by what they found.First, in 32 of the 45 invertebratespecies analyzed in detail, popula-tion abundances changed signifi-cantly. Second, animals that wereconsidered “southern” species werenow more common, with a con-comitant diminution in “northern”

species. On the other hand, twonorthern snails increased in abun-dance, but seemingly by migratingto a lower spot in the intertidal.Mean sea surface temperatures in-creased about 3/4oC, during these60 years, with an increase of ap-proximately 3o C during July throughAugust, the warmest months of theyear.

In Santa Cruz, John Pearse andhis students have continued to con-duct yearly censuses, and after 30years have only recently noticed arecovery in diversity at the site of thesewage outfall. Initially, at the outfallsite, coralline algae were the over-whelmingly dominant species. In thelate 1990s, signs of recovery at thissite, as measured by increased diver-sity, included the return of surfgrass,soft-bodied invertebrates such asbryozoans, tunicates, and sponges,and some of the delicate foliose redalgae. This is an excellent example ofa long-term study that has capturedimportant changes resulting from al-leviating pollution stress.

One important consequence ofthe Exxon Valdez spill was new fund-ing of large-scale, publicly and pri-vately funded censuses. These includeefforts by the National Marine Sanc-tuaries, the Partnership for Interdis-ciplinary Studies of Coastal Oceans(PISCO), the National Park Services(including an All-Taxa BiologicalInventory at Tomales Bay in north-ern California), and the Multi AgencyRocky Intertidal Network (MA-RINE), a consortium of agencies andinstitutions in southern California.

In some cases, innovative inter-agency alliances have been forgedbetween disparate groups, unified bythe desire to document and preserveour marine heritage. Fishermen, stu-dents, volunteers, and local residentshave been recruited to help univer-sity and government scientists withtheir census work. Traditional toolssuch as the plant press, transect, andquadrat are now augmented withGPS and GIS technology, and data-bases and images are often broadly

available on the Internet.We are now in the era of synthe-

sis and long-term, carefullydesigned experiments to try to teaseout the mechanism of change inthe intertidal zone. The need for anaccurate inventory of California’s in-tertidal flora is clear. With efforts ofvolunteers and professionals alike wewill hopefully be able to have a morecomplete appreciation for the diver-sity of this state’s seaweed flora, andfor the beauty of the balance withinand between intertidal ecosystems.


Barry,J.P, C.H. Baxter, R.D. Sagarin,and S.E. Gilman. 1995. Climate-related, long-term faunal changes ina California rocky intertidal com-munity. Science: 267:672-675.

Dawson, E.Y. and M. Foster. 1982(2nd Edition). Seashore Plants of Cali-fornia. UC Press.

Murray, S.N., J. Goodson, A. Gerrard,and T. Luas. 2001. Long-termchanges in rocky intertidal seaweedpopulations in urban southern Cali-fornia. J. Phycol. 37(3):37-38.

Pearse, J. Personal communication.Raimondi, P.T., R.F. Ambrose, J.M.

Engle, and M. Wilson. 1999. Moni-toring of rocky intertidal resourcesalong the central and southern Cali-fornia mainland. Three-year reportfor San Luis Obispo, Santa Barbara,and Orange Counties (Fall 1995,Spring 1998). Technical Report. USDepartment of Interior, MineralsManagement Service, Pacific OCSRegion (MMS Cooperative Agree-ment No. 14-35-0001-30761 withSouthern California Educational Ini-tiative, Marine Science Institute, Uni-versity of California, Santa Barbara).

Ricketts, E.F. 1992. Between PacificTides. Stanford University Press.Stanford, CA.

Thom, R. M. and Widdowson, T. B.1978. A resurvey of E. Yale Dawson’s42 intertidal transects on the south-ern California mainland after 15 years.Bull. S. Calif. Acad. Sci. 77:1-13.

Kathleen Dickey, PO Box 314, La Honda,CA 94020. [email protected]

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

If you are just beginning to learnabout the wonders of seaweeds, thefollowing list of reference books andwebsites will help you along your roadof discovery.

BOOKSBOOKSBOOKSBOOKSBOOKS

Abbott, I.A. and G.J. Hollenberg.1976. Marine Algae of California.Stanford University Press. Stanford,California, USA.

Dawson, E.Y. and M.S. Foster.1966. Seashore Plants of California.University of California Press. Berke-ley, California. Revised 1986.

Druehl, L.D. 2000. Pacific Seaweeds:A Guide to the Common Seaweeds of theWest Coast. Harbour Publishing. Ma-deira Park, British Columbia, Canada.

Lee, R.E. 1980. Phycology. Cam-bridge University Press. Cambridge,England.

Mondragon J. and J. Mondragon.2003. Seaweeds of the Pacific Coast: Com-mon Marine Algae from Alaska to BajaCalifornia. Sea Challengers. Monterey,CA.

Ricketts, E. F. et al. 1985. BetweenPacific Tides. Stanford UniversityPress. Stanford, CA, USA.

WEBSITESWEBSITESWEBSITESWEBSITESWEBSITES

Seaweed information andidentification

www.seaweed.ie. The National Uni-versity of Ireland, Galway. This com-prehensive website includes databaseand images of seaweed from aroundthe world.

www.mbari.org/~conn/botany/.Monterey Bay Aquarium ResearchInstitute, Monterey, California. Thissite includes student work from theHopkins Marine Laboratory MarineBotany class.

ucjeps.berkeley.edu/guide/. DeCew’sGuide to the Seaweeds of British Co-lumbia, Washington, Oregon, andNorthern California. See page 12.

www.coml.org/coml.htm. Census ofMarine Life. This is a global networkof researchers from more than 50countries engaged in a ten-year ini-tiative (begun in 2000), with the goalof assessing and explaining the diver-sity, distribution, and abundance ofmarine life in the oceans—past,present, and future. This endeavor isfunded by the Arthur P. Sloan Foun-dation. Two contributors to this siteare the Oceanic Biogeographic Infor-mation System(OBIS) and the Natu-ral Geography in Shore Areas(NaGISA).

Seaweed as food

www.seaweed.net. Mendocino SeaVegetable Company, MendocinoCounty, California.

www.loveseaweed.com. Rising TideSea Vegetable Company, MendocinoCounty, California.

Intertidal monitoring efforts

www.marine.gov. Multi AgencyRocky Intertidal Network. This site

represents a consortium of institu-tions, state and federal agencies, andprivate endeavors in southern Califor-nia conducting long-term intertidalmonitoring projects. Agencies includeCalifornia State University, Fullerton,Minerals Management Service, Uni-versity of Southern California, Uni-versity of California (Los Angeles, SanDiego, Santa Barbara, and Santa Cruzcampuses), Regional Water QualityControl Boards, PISCO (see websitebelow), California Department of Fishand Game, the United States Navy,the National Marine Sanctuary, andthe National Park Service.

www.bio logy.ucsd.edu/labs/roy/CBRISC/summary.html. Conservationand Biodiversity of the Rocky Inter-tidal of Southern California (with theacronym CBRISC, pronounced“SeaBrisk”). This site is aimed at assess-ing the health of Southern California’scoastal marine environment.

www.piscoweb.org. PISCO (Partner-ship for Interdisciplinary Studies ofCoastal Oceans). PISCO is a large-scale marine research program thatexamines near-shore marine systems.The program is based at four univer-sities along the Pacific coast, and isfunded by the Packard Foundation.

www.farallones.org/education_page/studentprojects. html#rocky. FarallonMarine Sanctuary Association(FMSA). Gulf of the FarallonesNational Marine Sanctuary, SanFrancisco, California Educators workwith FMSA on two intertidalmonitoring programs for students.

limpets.noaa.gov/. Long term moni-toring program and experiential train-ing for students (LIMPETS). Olym-pic Coast, Cordell Bank, Gulf of theFarallones, Monterey Bay, and Chan-nel Islands.

w w w. m b n m s - s i m o n . o r g / . T h eMonterey Bay National Marine Sanc-tuary (MBNMS). One of 13 Nationalmarine sanctuaries around the country,MBNMS has just posted an interactivedatabase at their website, called theSanctuary Integrated MonitoringNetwork (SiMoN).

SEAWEED RESOURCES

Compiled by Kathleen Dickey


The red alga, Phycodrys setchellii. Linedrawing, to be posted soon to the DeCewWebsite (see page 12), used withpermission from P. Silva.


NOTES AND COMMENTS

CNPS FELLOWS NOTCNPS FELLOWS NOTCNPS FELLOWS NOTCNPS FELLOWS NOTCNPS FELLOWS NOT

UNDER–APPRECIATEDUNDER–APPRECIATEDUNDER–APPRECIATEDUNDER–APPRECIATEDUNDER–APPRECIATED

The January 2003 issue of Fremontia(Vol. 31(1):27) listed California NativePlant Society Fellows, but some ofthose honored with the title were in-advertently omitted from this list.Apologies are given to these Fellows,their friends, and families. Although aFellow’s name might have been over-looked, certainly their participation inthe society is or has been greatly ap-preciated.

Fellows names are listed below withthe date the title was received.

Mary Ann Matthews, 1993Mary Rhyne, 1991Florence Marsh, 1991

Beatrice Howitt, 1973Jim Griffin, 1993Jake Sigg, 1997Joan Stewart, 1999Randall Morgan, 2000In addition, there may be others

who should be included on this list. Ifyou know if this is so and the date theybecame Fellows, please contact theEditor.

SUDDEN OAK DEATHSUDDEN OAK DEATHSUDDEN OAK DEATHSUDDEN OAK DEATHSUDDEN OAK DEATH

UPDATESUPDATESUPDATESUPDATESUPDATES

In April of 2004, wood rose (Rosagymnocarpa) was added to the list ofnearly 30 confirmed Phytophthoraramorum host plants. Wood rose is

native to California, and is commonlyfound in a wide range of habitats. Itis popular in the horticultural indus-try and is readily available from na-tive plant nurseries in California, Or-egon, Washington, and British Co-lumbia.

As a host plant for the Sudden OakDeath pathogen, wood rose falls un-der Phytophthora ramorum regulations,requiring inspection before being cer-tified for shipment from Californianurseries. For those planting nativesin your garden, be careful to obtainstock that is not infected with thispathogen.

For more information about Sud-den Oak Death, go to the CaliforniaOak Mortality Task Force (COMTF)website at: www.suddenoakdeath.org.

California state Fish and Gameregulations allows individuals to col-lect up to ten pounds of seaweed fornoncommercial uses. The best time tocollect is just after a storm or whenthe tide is going out, in order to en-sure freshness. The best places to col-lect drift seaweed are open oceanbeaches and rocky shores, away fromcreeks or rivers. Seaweeds from baysand inlets in California are best leftalone. There are no poisonousseaweeds, although one genus,Desmarestia, contains hydrosulfuricacid and is quite acrid. Others, suchas sea lettuce (Ulva), are just too toughto eat.

GENERAL RULESGENERAL RULESGENERAL RULESGENERAL RULESGENERAL RULES

1) Before you pick any seaweed,make sure that there are many moreplants of the same species nearby. Forexample, in the case of the sea palm(Postelsia), one study shows that thereneed to be at least 120 plants in a standfor it to reproduce.

HARVESTING AND COLLECTING SEAWEEDS

2) Some seaweeds are quite rare,so if you find an individual plant ofany species, leave it alone.

3) Learn about the reproductivecycle of the seaweeds you are collect-ing, and what a fertile plant lookslike. By not collecting fertile plants,the population will be able to replen-ish itself.

4) Do not assume that because aplant is growing in the ocean it will betoxin-free. Some seaweeds selectivelyuptake and concentrate minerals. Donot collect seaweed near a city or neara stream or river that might carry ag-ricultural runoff into the ocean. As ageneral rule, bay waters (especially SanFrancisco Bay) should be off-limits.


Dawson, E.Y. and M. Foster. 1982. Sea-shore Plants of California. Universityof California Press. Berkeley, CA.

Druehl, L. 2001. Pacific Seaweeds.Harbour Publishing. Madeira Park,BC Canada.

The red alga, Ceramium californicum.Line drawing, to be posted soon to theDeCew Website (see page 12), used withpermission from P. Silva.

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

BOOK REVIEW

Pacific Seaweeds: A Guide to theCommon Seaweeds of the WestCoast, by Louis Druehl. 2000.Harbour Press, Madeira Park, BritishColumbia, Canada. Softcover, $17.95.

Louis Druehl has been studying andteaching about seaweeds for more than30 years, and recently started farmingkelps in British Columbia. In this bookhe presents information and fascinat-ing lore about seaweeds aimed at ageneral, interested audience. He in-

corporates taxonomy based on exter-nal morphology (for example, crusts,globular, small flat blades, cylinders)with a small “thumbnail” dichoto-mous key. The photographs and linedrawings are very helpful in the field.As one must in any small book,Druehl focuses on some 100 of themore common seaweeds, perhapswith a disproportionate emphasis onhis specialty, the “browns.”

My favorite parts of this book are

the fascinating stories and asidesthat range from the intriguing tothe preposterous. For example, he de-scribes the sea palm, Postelsiapalmataeformia, as being “severelyattached to a rock.” Additionally, heincludes many simple recipes, somecomplete with verse. Altogether, thisis a satisfying introductory book toaccompany you in forays to thetidepools. It includes a helpful glos-sary and list of references.

RECIPE FOR MARINATED RECIPE FOR MARINATED RECIPE FOR MARINATED RECIPE FOR MARINATED RECIPE FOR MARINATED ALARIAALARIAALARIAALARIAALARIA

Thanks to Louis Druehl and Rae Hopkins,Canadian Kelp Resources, Ltd.

Ingredients1 cup dried Alaria, cut in strips (use scissors)1/2 cup rice vinegar3 Tbsp soy sauce1 Tbsp sugar1 tsp salt

Optional and delicious additionsThinly sliced cucumberFinely grated carrotsSliced green onions

InstructionsPlace Alaria strips in bowl and cover with warm water for 1/2 hour to rehydrate. Drain. In a small bowl orjar, mix the rice vinegar, soy sauce, sugar and salt, and mix well. Pour over Alaria. Serve with wheat crackersas an appetizer. To serve as a salad, add thinly-sliced cucumbers and serve on leaves of butter lettuce. Garnish with slicedgreen onions.


Weeds and Wildflowers(Volume 31, No. 2)

I wanted to say how much I enjoyedJake Sigg’s article on weeds, and alsothe article by Marjorie Schmidt! Itwould be great if Fremontia ran an ar-ticle of historical quotes in every is-sue. Jake’s article was so well-writtenand thought-provoking—I had not

LETTERS TO THE EDITOR

considered the effect of plaintain andweeds like it before.

Kathy KramerEast Bay Chapter

Weed Reference Correction(Volume 31, No. 4:34)

In the “Selected Articles on Weeds,”the reference to “Control of the

Aleins” should be ascribed to DorisAnn Hoover, who sadly passed awayrecently. Doris was one of the origi-nal weed warriors in the LA Chapter,and these comments appeared first, ifI remember correctly, in the localToyon.

Michael O’BrienLos Angeles/Santa Monica

Chapter

The kelp, Alaria marginata. Linedrawing from DeCew Guide, usedwith permission from P. Silva.

V O L U M E 3 2 : 2 , A P R I L 2 0 0 4

CLASSIFIED ADS

Classified ad rate: $1.00 per word,minimum $15; payment in advance.Address advertising inquiries and copy to:CNPS, 2707 K Street, Suite 1, Sacra-mento, CA 95816-5113. (916) 447-2677or fax (916) 447-2727.

PUBLICATIONS

Flora & Fauna Books, 121 First AvenueSouth, Seattle WA 98104, Tel. (206)623-4727, Fax (206) 623-2001, [email protected], Specializing in Botany,Gardening, Birding, and Ecology, bothnew and out-of-print. We carry a largeinventory of floras, keys, and field guidesfor the west coast and worldwide. Alarge selection of our inventory is nowavailable on the web: www.abebooks.com/home/FFBOOK/.

ART

Notecards, Prints, and Originals. Visitwww.VorobikBotanicArt.com. PO Box866, Lopez Island, WA 98261

Botanical prints, note cards, postcards,Plants of the Lewis and Clark Expedition

notecards, books, fruit crate labels, handtowels, and T-shirts focusing onCalifornia native plants from SierraNature Prints. Animal puppets, too. Visitwww.sierranatureprints.com

NURSERIES AND SEEDS

Telos Rare Bulbs. Bulbs for your gar-den, restoration projects, landscaping.Many Calif. native species, includingCalochortus, Fritillaria, Brodiaea rela-tives, Erythronium. Catalog $3.00. Freeshipping in USA. P.O. Box 4978, Arcata,CA 95518. www.telosrarebulbs.com.

ECOLOGICAL RESTORATION

Native California grassland and desertecological restoration standards andcosts, plus pictures showing results.www.ecoseeds. com/standards.html.

SERVICES

Nature landscape design. LandscapeDesign that celebrates the rich heritageof California’s native flora. Duber Land-scape Design, CA license #4316. (510)524-8665.

MATERIALS FORPUBLICATION

Members and others are in-vited to submit material for pub-lication in Fremontia. Instruc-tions for contributors can befound on the CNPS website,www.cnps.org, or can be re-quested from Fremontia Editor,Linda Ann Vorobik, [email protected], or c/o University andJepson Herbaria, 1001 ValleyLife Sciences Bldg. #2465, Uni-versity of California, Berkeley,CA 94720-2465.

FREMONTIA EDITORIALADVISORY BOARD

Ann Bradley, Susan D’Alcamo-Potter, Ellen Dean, KathleenDickey, Phyllis M. Faber, BartO’Brien, John Sawyer, JimShevock, Teresa Sholars, NevinSmith, Dieter Wilken, JohnWilloughby, Darrell Wright

❏ Enclosed is a check made payable to CNPS Membership Gift:

❏ Charge my gift to ❏ Mastercard ❏ Visa Added donation of:

Card Number TOTAL ENCLOSED:

Exp. date

Signature

Phone

Email

CNPS member gifts allows us to promote and protect California’s native plants andtheir habitats. Gifts are tax-deductible minus the $12 of the total gift which goestoward publication of Fremontia and the CNPS Bulletin.

Please Join Today!

❏ $20 Limited Income ❏ $35 Individual ❏ $45 Family/International ❏ $75 Supporting❏ $100 Plant Lover ❏ $250 Patron ❏ $500 Benefactor ❏ $1,000 Mariposa Lily

Please make your check payable to “CNPS” and send to: California Native Plant Society, 2707 K Street, Suite 1,Sacramento, CA 95816-5113. Phone (916) 447-2677; fax (916) 447-2727; www.cnps.org.

❏ Enclosed is a matching gift form from my employer

❏ I would like information on planned giving

NAME

ADDRESS

CITY STATE ZIP

V O L U M E 3 2 : 1 , J A N U A R Y 2 0 0 4

California N

ative Plant S

ociety2707 K

Street, S

uite 1S

acramento, C

A 95816-5113

Address S

ervice Requested

Paul C. Silva and Kathy Ann Miller,have through their combined effortsoffered several perspectives fromwhich to consider algae. Silva relatesstories of their discovery and how theyare harvested for useful products. Healso carefully describes how currentsalong our coast contribute to the greatdiversity of California seaweeds.Miller informs us of invasive seaweedsthat we, as stewards to natural envi-ronments, should be on the lookoutfor. Dickey provides background in-formation on seaweed research as wellas how seaweeds are monitored, andwhy it is important to continue study-ing them. And if you are inspired tolearn more, look to the back pages fora list of Web and other resources that

Dickey has provided, or try the recipefrom the book review on page 32.

This issue begins my fourth year asEditor of Fremontia, and during thattime my goal has been to provide avariety of articles that are backed byacademic research and yet are inter-esting and available to a broad audi-ence—with articles ranging from wild-flower appreciation and horticulture,through ecology and taxonomy, toadvocacy. I wish to make these issuesthe best they can be for our society,and welcome all letters that respondto individual articles or to the generalscope of our California Native PlantSociety journal, Fremontia.

Linda Ann Vorobik,Editor

FROM THE EDITOR

CONTRIBUTORS

Tom DeCew received his MS at Humboldt State Universityand his PhD at UC Berkeley. During his studies he canvassedWest Coast herbaria to compile phenology and distributiondata and searched the macroalgal literature for studies pertain-ing to West Coast seaweeds. The fruits of his labor resulted in“The Book,” a bibliography to the biology of common north-east Pacific seaweeds with line drawings and tabular displays oflatitudinal distribution and reproductive phenology. It was in-tended to complement Marine Algae of California by I.A. Abbottand G.J. Hollenberg. DeCew died prematurely in 1997, be-fore his Magnum Opus could be published.

Kathleen Dickey is a research associate at the herbarium, UCBerkeley. She received her MS at UCSC. She has worked as aconsultant for NOAA for the past four years, conducting inter-tidal censuses on Southeast Farallon Island, near San Francisco,California. Her particular area of interest is the pattern of dis-tribution of intertidal algae. She is currently designing a set of“seaweed flashcards” to help identify seaweeds in the field.

Kathy Ann Miller is the resident scientist at the University ofSouthern California’s Wrigley Marine Science Center on SantaCatalina Island. She earned her BA and PhD degrees in Botanyat the University of California at Berkeley. She specializes inseaweeds of California and adjacent coastlines. Her current re-search includes studies of the systematics, ecology, and evolu-tion of seaweed and eel grass populations around the Califor-nia Channel Islands.

Paul C. Silva is a native son of California. He received his BAat the University of Southern California, his MA at StanfordUniversity, and his PhD at the University of California, Berke-ley. He has been on the staff of the University Herbarium since1960 and is considered a world authority on seaweeds.

T he seashore provides a con- trasting experience to thatwhich we find with walks to

meadow or mountain. The smell ofsalt, view of ocean, and sound of windand wave all combine to make a visitto the intertidal unique. This specialissue of Fremontia leads us into theworld of seaweeds, and provides pre-liminary information along with a fewtools of understanding that may makeyour next visit there as joyful as anymeadow stroll. And if you glory in di-versity and detail, you will be delightedby this group of plants that differ sofrom our more familiar flowers offorest and field.

The Convening Editor, KathleenDickey, along with the contributors

Nonprofit O

rg.

U.S

. Postage

PA

IDO

akland, CA

Perm

it # 3729

Vol. 32, No. 1 January 2004 FREMONTIAvolume 32:1, january 2004 a journal of the california native...

Documents

Transcript of Vol. 32, No. 1 January 2004 FREMONTIAvolume 32:1, january 2004 a journal of the california native...