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Vol. 32, No. 1 January 2004 FREMONTIA

A JOURNAL OF THE CALIFORNIA NATIVE PLANT SOCIETY

IN THIS ISSUE:

SEAWEEDS TELL THEIR STORIES by Paul C. Silva 3

CALIFORNIA’S NON–NATIVE SEAWEEDS by Kathy Ann Miller 10

CALIFORNIA SEAWEED INDUSTRY by Paul C. Silva 16

DISTRIBUTION OF CALIFORNIA SEAWEEDS by Paul C. Silva 18

INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS by Kathleen Dickey 27 VOLUME 32:1, JANUARY 2004 SPECIAL ISSUE: SEAWEEDS CALIFORNIA NATIVE PLANT SOCIETY

FREMONTIA CNPS, 2707 K Street, Suite 1; Sacramento, CA 95816-5113 (916) 447-CNPS (2677) Fax: (916) 447-2727 VOL. 32, NO. 1, JANUARY 2004 [email protected]

Copyright © 2004 MEMBERSHIP California Native Plant Society Membership form located on inside back cover; dues include subscriptions to Fremontia and the Bulletin Distributed in June 2004 Mariposa Lily ...... $1,000 Supporting ...... $75 Linda Ann Vorobik, Editor Benefactor ...... $500 Family, Group, International . . . $45 Katheen Dickey, Convening Editor Patron ...... $250 Individual or Library ...... $35 Bob Hass, Proofreader Plant Lover ...... $100 Student/Retired/Limited Income . $20 Susan Cotterel, Designer STAFF CHAPTER COUNCIL CALIFORNIA NATIVE Sacramento Office: Alta Peak (Tulare) . . . . Joan Stewart PLANT SOCIETY Executive Director . . . . Pamela C. Bristlecone (Inyo-Mono) ...... Muick, PhD Sherryl Taylor Channel Islands ...... Lynne Kada Development Director . . . Michael Dedicated to the Preservation of Dorothy King Young (Mendocino/ Tomlinson the California Native Flora Sonoma Coast) . . . . Jon Thompson Membership Coordinator . . Marin The California Native Plant Society East Bay ...... Tony Morosco Lemieux El Dorado ...... Amy Hoffman (CNPS) is a statewide nonprofit orga- Sales Manager ...... Paul Maas Kern County . . . . . Laura Stockton nization dedicated to increasing the Bookkeeper . . . . Lois Cunningham Los Angeles/Santa Monica Mtns . . . understanding and appreciation of Betsey Landis California’s native plants, and to pre- At Large: Marin County ...... Bob Soost serving them and their natural habi- Fremontia Editor . . . . . Linda Ann Milo Baker (Sonoma County) . . . . . tats for future generations. Vorobik, PhD Reny Parker CNPS carries out its mission Sr. Policy Analyst ...... Emily Mojave Desert ...... Tim Thomas through science, conservation advo- Monterey Bay ...... Roberson, PhD cacy, education, and horticulture at Julie Anne Delgado So. California Regional Botanist . . . the local, state, and federal levels. It Mount Lassen ...... Jim Bishop Ileene Anderson monitors rare and endangered plants Napa Valley ...... Marcie Danner Interim Rare Plant Botanist . . . and habitats; acts to save endangered North Coast ...... Larry Levine Misa Ward areas through publicity, persuasion, North San Joaquin . . . . Gail Clark Vegetation Ecologist . . Julie Evens Orange County . . . . . Sarah Jayne and on occasion, legal action; pro- Redbud (Grass Valley/Auburn) . . . . vides expert testimony to government Vegetation Ecologist . . Anne Klein San Bruno Mtn. Project Coordinator Richard Hanes bodies; supports the establishment of Riverside/San Bernardino counties . . native plant preserves; sponsors work- Joe Cannon Legislative Advocate .Vern Goehring Katie Barrows days to remove invasive plants; and Sacramento Valley . . Diana Hickson Legal Advisor . . . . Sandy McCoy offers a range of educational activities San Diego ...... Dave Flietner including speaker programs, field trips, Website Coordinator ...... San Gabriel Mtns . . . . Lyn McAfee native plant sales, horticultural work- John Donaghue San Luis Obispo . . . . Dirk Walters shops, and demonstration gardens. Bulletin Editor . Michael Tomlinson Sanhedrin (Ukiah) . Chuck Williams Since its founding in 1965, the tra- Santa Clara Valley . . Georgia Stigall ditional strength of CNPS has been Santa Cruz County . Janell Hillman BOARD OF DIRECTORS its dedicated volunteers. CNPS ac- Sequoia (Fresno) . . . . Warren Shaw tivities are organized at the local chap- Carol Baird, Jim Bishop, Vern Shasta ...... Dave DuBose Sierra Foothills (Tuolumne, Cala- ter level where members’ varied in- Goerhing, Steve Hartman, Diana veras, Mariposa) . . . . Patrick Stone Hickson, Lynn Houser, Lynne Kada, terests influence what is done. Volun- South Coast (Palos Verdes) ...... teers from the 33 CNPS chapters an- David L. Magney, Sandy McCoy, J. Barbara Sattler nually contribute in excess of 87,000 Spence McIntyre, Carol Witham Tahoe ...... Michael Hogan hours (equivalent to 42 full-time em- (President) Willis L. Jepson (Solano) ...... ployees). Allison Fleck CNPS membership is open to all. PROGRAM DIRECTORS Yerba Buena (San Francisco) . . . . . Members receive the quarterly jour- Mark Heath nal, Fremontia, the quarterly statewide CNPS Press ...... Holly Forbes and Gail Milliken MATERIALS FOR Bulletin, and newsletters from their PUBLICATION local CNPS chapter. Conservation. . . . . David Chipping Horticulture. . . Peigi Duvall CNPS members and others are wel- Fremontia logo (by L.A. Vorobik) re- Posters ...... Bertha McKinley come to contribute materials for pub- printed from The Jepson Manual, J. and Wilma Follette lication in Fremontia. See the inside Hickman, Ed., 1993, with permission Rare Plants...... Ann Howald back cover for manuscript submission fromthe Jepson Herbarium, UC. © Re- Vegetation . . . . Todd Keeler-Wolf instructions. gents of the University of California. Prepress by ScanArt / Printed by Craftsman Trade Press

VOLUME 32:1, JANUARY 2004 CONTENTS GUEST EDITORIAL ...... 2

SEAWEEDS TELL THEIR STORIES by Paul C. Silva ...... 3 For 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 Silva relates the tales of their discovery and naming by Russian and Swedish explorers, keen-eyed Victorians, and resourceful Midwesterners.

CALIFORNIA’S NON–NATIVE SEAWEEDS by Kathy Ann Miller...... 10 It is well known that our terrestrial flora has numerous introduced species, some of which are especially 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 INDUSTRY by Paul C. Silva ...... 16 Many of us are familiar with the red alga “nori,” which wraps around the rice in sushi. Few of 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 SEAWEEDS by Paul C. Silva...... 18 Seaweeds are restricted vertically by the availability of light in deep water and by their ability to withstand desiccation in the intertidal zone. Each species has an optimal temperature range which 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 SEAWEEDS by Kathleen Dickey ...... 27 Marine biologists have documented the California intertidal for more than a century, but there remains much to decipher. Threats to the intertidal, both natural and man-made, have intensified efforts to monitor and further understand this fascinating ecosystem.

SEAWEED RESOURCES ...... 30 HARVESTING AND COLLECTING SEAWEEDS ...... 31 NOTES AND COMMENTS ...... 31 LETTERS TO THE EDITOR ...... 32 BOOK REVIEW ...... 32

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 life in central and southern California nearshore communities. Photograph courtesy of J. and J. Mondragon (www.mondragonphotos.com).

VOLUME 32:1, JANUARY 2004 FREMONTIA 1 GUEST EDITORIAL USEFUL WEBSITES AND CONTACT ur world is both challenging and inspiring, with moments of urgency INFORMATION O juxtaposed with those of riveting beauty. The world of seaweeds is no exception. Last fall there was news of yet another old oil tanker off the coast of Seaweed websites: Spain that was in danger of breaking and spilling its cargo. To add insult to See “Seaweed Resources,” p. 30 injury, this occurred along the same stretch of coast that had been inundated California Native Plant with oil just the year before. A request went out around the world via the Algae Society (CNPS): Listserv (www.seaweed.ie) for baseline data and for assistance with tackling the www.cnps.org, with links to formidable task of remediation. Despite continuous wake-up calls, we still seem conservation issues, chapters, to be “asleep at the wheel” with respect to protecting the intertidal regions of publications, policy, etc. the world and preventing these disasters from recurring. To sign up for “NPCC News,” Fortunately, the scientific community has been “waking up” for nearly half e-mail news on native plant a century and is beginning to compile and synthesize long-term data sets that science and conservation, send will document changes due to such events, and will thus provide ammunition in a request to [email protected]. the fight for laws and policies that will protect our coasts and their native ecosystems, including the seaweed flora. For updates on conservation Worldwide this flora is impressive: there are some 8,000 species of marine issues: algae—simple, salt-water dwelling organisms that, to quote Dr. Mike Guiry, Audubon Society www.audubon.org professor of phycology, “fall into the rather outdated general category of ‘plants.’ ” Center for Biological Diversity The algae include three major groups which have evolved quite separately and are www.sw-center.org described by their accessory pigments: the reds (the most abundant), the browns Natural Resources Defense (which include the kelps), and the greens (which may actually be part of the Plant Council Kingdom). www.nrdc.org Along the coast of California we have a particularly rich flora of seaweeds, Sierra Club which includes some 700 species. This great diversity is due in part to the cold, www.sierraclub.org upwelling ocean waters that bring nutrients and oxygen to our coastal intertidal Wilderness Society regions. In addition, the geological forces that have formed our coast resulted www.wilderness.org in a gorgeous patchwork of uplifted rock, providing a multitudinous array of substrates for the seaweeds to anchor themselves with their holdfasts. Read For voting information: about the diversity of California seaweeds in Paul Silva’s article on biogeogra- League of Women Voters phy on page 18. And, as in the terrestrial flora, some seaweeds found in www.lwv.org, includes online voter California are natives of other regions. Kathy Ann Miller writes about weedy guide with state-specific nonparti- seaweeds in her article on page ten. san election and candidate information. Seaweeds themselves boast some amazing records. Consider the following: They have been found as deep as 268 meters in the waters of the Caribbean, as US Senate www.senate.gov well as under the ice at Antarctica. The giant kelp, Macrocystis pyrifera, can grow up to two feet per day. Recently a whole new group of red coralline seaweeds US House of Representatives have been discovered called rhodoliths, individuals of which are at least 150 www.house.gov years old, or which, if you take into account their capacity to reproduce California State Senate vegetatively, may be thousands of years old. www.sen.ca.gov Whether we are aware of it or not, our lives are impacted daily by marine California State Assembly algae, as they are used commercially in many familiar foods and products, such www.assembly.ca.gov as the thickening agent of toothpaste and paint, and as the emulsifier in To write letters: chocolate milk. They are the basis of a thriving industry off the southern California coast. Paul Silva writes of seaweeds and industry in his article on President George W. Bush The White House page 16. 1600 Pennsylvania Ave. NW Most native plant enthusiasts will celebrate with curiosity and appreciation Washington, DC 20500 California’s intertidal flora, but for those of you who might have less interest in Senator Barbara Boxer these salt-water dwellers, you are at first teased in with two stories of seaweed or Senator Dianne Feinstein discovery (page three) by Paul Silva. Professor Silva deserves special US Senate acknowledgement for his contribution to this issue, as he is one of the foremost Washington, DC 20510 authorities on California seaweeds. Your CA Representative —Kathleen Dickey US House of Representatives CNPS Santa Clara Valley Chapter Washington, DC 20515 Member of the Fremontia Editorial Advisory Board

2 FREMONTIA VOLUME 32:1, JANUARY 2004 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 diverse and beautiful plants. Here are two such stories.

SEA PALM

The western edge of the Ameri- can continent was put on the map, literally and figuratively, when, on the 15th of July, 1741, the St. Paul, commanded by Alexei Chirikov, reached the shore of an island in what today we know as the Alexander Archipelago in southeast- ern Alaska. Vitus Bering, heading the Second Kamchatka Expedition and commanding the St. Peter, made landfall at Cape St. Elias at the tip of Kayak Island a few days later (on July 18th). The two ships had pro- ceeded independently after having lost sight of one another in the fog. The St. Paul returned to Kamchatka on October 8th, with most of the crew dead from scurvy. The Saint Peter was undergoing even greater hardships, being wrecked on the westernmost side of the Commander Islands close to Kamchatka. The crew overwintered on this island, which they named Bering Island in memory of their captain, who had died shortly after the shipwreck. In the spring, enough boards and nails were salvaged from the wreck to construct a small boat, which enabled only a handful of survivors to return to Kamchatka on August 25, 1742. Tales of the great wealth of natural resources of “Bolshaya Zemlya” (“Big Land”), as the Rus- sians initially called Alaska, quickly resulted in a flood of Russian ad- venturers, mostly trappers and fur traders. Their brutal treatment of Postelsia palmaeformis, the sea palm, was named after the Estonian geologist and artist the Aleut people and the near-ex- who painted seaweeds from nature during the 1826-1829 round-the-world voyage on the vessel Seniavin. Illustration from F.J. Ruprecht, 1852: Neue oder unvollständig tinction of the sea otter are well- bekannte Pflanzen aus dem nördlichen Theile des stillen Oceans [New or incompletely known dark episodes of history. By known plants from the northern parts of the Pacific Ocean], St. Petersburg.

VOLUME 32:1, JANUARY 2004 FREMONTIA 3 selling the fort and its contents to book, Illustrationes Algarum, con- John Sutter. Vosnesensky by this tains the original descriptions of time was collecting in Baja Califor- many species that are common in nia. His collections were sent to the California, although no collections Academy of Sciences, located in St. were made south of southeastern Petersburg. Alaska. The specific epithet, The seaweeds collected by palmaeformis, is self-explanatory, the Vosnesensky were studied by Franz resilient plants bending over in the Josef Ruprecht, who was born in surf reminiscent of palm trees dur- Austro-Hungary, studied medicine ing a hurricane or typhoon. Postelsia palmaeformis, the sea palm, in Prague, and became the curator It may or may not be surprising growing with red algae and gooseneck of botany at the Academy of Sci- that none of the several botanists barnacles. Photograph by G. Hansen. ences at St. Petersburg in 1839. In who sampled the marine flora of 1852 Ruprecht published a paper in Monterey during the late 18th cen- which he treated only five seaweeds tury and early 19th century collected 1792 there was sufficient Russian and one seagrass from Sonoma this quintessentially unique alga. presence in Alaska to warrant the County, but which is justly famous Perhaps the reason for this lacuna establishment of a permament not only for the three new genera is that the sea palm, hanging on for settlement on Kodiak Island, fol- that he described, but also for the dear life on wave-swept rocks, sig- lowed in 1799 by the founding of oversized and remarkably lifelike nals would-be collectors to be on the town of Novoarkhangelsk (New lithographs. The plate for Egregia guard lest they find themselves in a Arkhangel, now known as Sitka). menziesii is 1.46 meters long, has similar life-threatening position. In order to supply food for the eight folds, and is spliced only once. Vosnesensky provided ample Russian colony, the government, in One of Ruprecht’s new genera collection data. The three speci- 1812, secured an area on the was the sea palm, which he named mens illustrated in Ruprecht’s plate Sonoma County coast, which was Postelsia palmaeformis. The generic were collected on an islet at the planted with crops and protected name honors Alexander Philipou entrance to Bodega Bay, called by by a fortification. It was named Postels, an Estonian-born geologist the natives “Omújüpai.” They called Rossiya (later called Fort Ross by and artist who had painted seaweeds the sea palm “Kakgunu-chale,” Americans) while a nearby river was from nature while on board the whose meaning I do not know, but named Slavianka (later called Rus- Seniavin during the voyage around certainly it would have nothing to sian River by Americans). Trappers the world commanded by Friedrich do with palm trees. and fur traders were soon raiding Lütke (also transliterated as Litke) The sea palm belongs to the the coast as far south as San Fran- in the period from 1826 to 1829. In order Laminariales of the class cisco. Russians competed with 1840, Postels and Ruprecht together Phaeophyceae (brown algae). It is Americans for the eggs of sea birds, had published a monumental “el- one of several very distinctive kinds another lucrative item of trade. By ephant” folio volume treating the of kelp, as members of this order 1840 nature’s bounties were seri- collections of seaweeds made dur- are called. It is essentially an an- ously depleted in northern Califor- ing that expedition. This very rare nual, growing rapidly in spring, nia, and the Russian government, feeling the hot breath of the British to the north and Mexicans to the south, decided to abandon the project. The prestigious Academy of Sciences in St. Petersburg, upon hearing this decision, arranged for a young zoologist and naturalist, Ilya Vosnesensky, to go to Rossiya and survey the nearby animals and plants. The Sonoma coast has many spectacular kelps, and these were collected by Vosnesensky, who ig- nored the smaller seaweeds. His In 1852 Ruprecht described one species of the surfgrass genus Phyllospadix mission completed, the Russian gov- (Zosteraceae Family). This plant is one of the few angiosperms that has adapted to ernment closed Rossiya in 1842, life in the intertidal surf. Photograph of flowers and leaves by S. Sharnoff.

4 FREMONTIA VOLUME 32:1, JANUARY 2004 maturing in late summer, discharg- ous generation. the expedition, C.A. Virgin. The ing spores, and remaining until the At the same time that Ruprecht epithet, palmaeformis, again is self- waves destroy it by relentless pound- described Postelsia, a Swedish explanatory. Areschoug’s paper was ing or knock it off its perch. In the around-the-world expedition on published in 1853, only a year after intense competition for space in the board the frigate Eugenie stopped Ruprecht published Postelsia. intertidal zone, Postelsia often finds in San Francisco Bay, where the itself perched on mussels or bar- naturalist, Nils Johan Andersson, nacles, which may be swept off the collected a specimen of sea palm THE EPHEMERAL rocks all together. Overwintering is (probably from Point Lobos or LAMINARIA accommodated by the microscopic Lands End on the Pacific side of sexual plants that develop from the San Francisco). This specimen was Our second story, involving an- discharged spores. The fertilized referred to a Swedish botanist, John other distinctive kelp, begins on the egg begins development of the erect Erhart Areschoug, who specialized Monterey Peninsula, nearly a half- plant, but then waits until early in the study of kelps. Unaware of century later. Laminaria ephemera, spring, when rapid growth occurs. Ruprecht’s work, Areschoug de- as the specific epithet implies, is The spores often stick to the sub- scribed the new species as Virginia ephemeral, appearing in March and strate close to the mother plant, so palma-maris. The generic name may disappearing in August. It is at- that in early spring a grove of very bring to mind the State of Virginia tached to the substrate by means of small “palm trees” may be seen sur- or perhaps the Virgin Mary, but in a holdfast, from which there arises rounding the remains of the previ- fact it commemorates the leader of a stem ending in a simple blade. The blade produces an abundance of spores, which settle on a sub- strate and germinate to produce minute sexual plants. After an egg has been fertilized by a motile male gamete, the resulting zygote re- mains dormant or undergoes only the first stages of development into a macroscopic plant during winter. Growth of the new plants is very rapid in early spring, followed by maturation, spore discharge, and disintegration. Laminaria ephemera is one of only a few species of the genus that are annual. In most spe- cies the macroscopic plant is peren- nial, the stem continuing to increase in diameter while the blade is re- newed at the beginning of each growing season after being torn away by the waves. Laminaria ephemera is not only ephemeral, but also elusive, appear- ing in large numbers at only a few widely separated sites. It thus es- caped the notice of naturalists on early expeditions. It did not escape the sharp eyes of Mrs. J.M. Weeks, however, who first recognized its distinctness dur- ing one of her frequent forays to Carmel Bay. Mrs. Weeks was an enthusiastic and dedicated seaweed The life cycle of the bull kelp (Nereocystis) is typical of all kelps, including Postelsia palmaeformis and Laminaria ephemera. Illustration by L. Scharf, from Seaweeds of the collector, as were several other la- Pacific Coast by J. Mondragon & J. Mondragon (Sea Challengers 2003). dies living in Pacific Grove during

VOLUME 32:1, JANUARY 2004 FREMONTIA 5 Pressed specimens of Laminaria epehemera collected by Mrs. J.M. Weeks, whose persistence and careful collecting habits succeeded in getting Professor W.A. Setchell’s attention. Image use courtesy of the University Herbarium, UC Berkeley.

6 FREMONTIA VOLUME 32:1, JANUARY 2004 the 1890s. Some were devoted to enough for a set.” (The “set” refers making spectacular mounts on to the Phycotheca Boreali-Ameri- deckle-edged cardboard to be sold cana, a set of dried specimens sold in curio shops, perhaps to end up to subscribers.) The remainder of on someone’s mantle in San Jose. the letter deals with possible ar- Mrs. Weeks had a keen scientific rangements whereby Setchell could interest and rejoiced when she either go to see the Laminaria or learned that the University of Cali- have it collected for him by a resi- fornia had added a phycologist to dent of Carmel. A year later, in a its faculty. That man was William letter from “Carmel City” dated A. Setchell, who was brought to April 13, 1898, Mrs. Weeks wrote: Berkeley in 1895 to be the head of a “The Laminaria I sent you last newly organized botany depart- spring is quite abundant. Do you ment. Setchell brought with him care to see any more of it? . . . The extensive knowledge of the algae, plants are older than they were last having studied under the tutelage year at this time. The stripes seem of two eminent cryptogamic bota- to be only an occasional feature / nists, Daniel C. Eaton of Yale Uni- The sporangia cover a large por- versity for his master’s degree, and tion of the frond. I noticed today William G. Farlow of Harvard Uni- on some I gathered to send away versity for his doctoral degree. With the disk was smooth & round about this background, he satisfied both a fourth of an inch in diameter . . . I factions on the faculty—Yalees and wish you might see them / They Harvardmen. seem of the Neriocystus [i.e., Arriving in Berkeley two months Nereocystis] type . . .”A few days later after classes had begun, Setchell was (April 24), following Setchell’s re- Professor W.A. Setchell, photographed impatient to sample the local flora. ply, Mrs. Weeks wrote: “In regard in his office, circa 1934. Photograph by During Christmas break, he to the Laminaria I sent you: I need W. Matthews, courtesy of the University boarded a coastal steamer bound not say it is a painful disappoint- Herbarium, UC Berkeley. for San Pedro with a stop in both ment to me not to be able to get at directions at Port Harford (now the plant. The portions of the beach dated June 1, 1899, she wrote: “The called Port San Luis). He collected that had enough of it accessable are Laminaria I sent is not in any par- seaweeds at the two sites. Shortly bedded in with sand and all but the ticular like the one we have been in after the Christmas break, Setchell tips are buried. The sand shifts in a the habit of calling L. Andersonia / received an introductory letter from day. I can see it out in the deeper It is not the color, thickness, nor Mrs. Weeks (dated February), water but as the surf is bad there I period of growth, or plan of form- which he promptly answered, and don’t feel strong enough to venture ing sporangia of that plant . . . I thus began a lively correspondence . . . I have been to a considerable thought when I commenced gath- that lasted through 1905. expense in coming down here & am ering it was young Neriocystus and The plant that was to be named not very comfortable in the work wondered it grew so long without Laminaria ephemera was first men- . . . But I was so interested in that an air bladder & when I found a tioned by Mrs. Weeks in a letter Laminaria that I came. Why do you fully developed plant with several from Pacific Grove dated April 22, call it Andersonii [Laminaria dark, even bands of different widths 1897, written upon her return from andersonii, a name misapplied to the [sporangial patches] running down an excursion to Carmel Bay made species correctly called Laminaria the length of it, it dawned upon me in a phaeton pulled by “Old Lottie”: setchellii]? It has no resemblance to that I had found a stranger to my “By this morns Express sent a can that plant. Its holdfast is different previous collection . . . I showed containing a Laminaria of which I from that, or indeed any other Lami- one of my mounts to Dr. Anderson once found, some three or four years naria I have known / Its stipe is [a physician in Santa Cruz who was a single specimen / I had begun to tender / Its blade similar to the keenly interested in seaweeds] & he think it a myth, as I could find no Neriocystus in substance and color.” asked if I had painted the stripes / more of it / Yestermorning I came Mrs. Weeks persisted, impa- He had never seen it. I was very upon a great amount of it but my tiently waiting for Setchell to study much in hopes you would get down containing vessels & little cart were and describe this interesting sea- in time to see & study it where it already full / I could not secure weed. In a letter from Santa Cruz grew, for I think it is short lived”.

VOLUME 32:1, JANUARY 2004 FREMONTIA 7 LEARNED AMATEUR CONTRIBUTES TO SEAWEED SCIENCE

Letter from Mrs. Weeks to Professor Setchell, demonstrating that although Mrs. Weeks was not a professional botanist, she was quite 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 from Santa Cruz & the flat thin bladed Pelvetia that I sent from there, were they examined and what are they? Did you find more than one oosphore in the flat Pelvetia[?] There is a plant growing abundantly in Carmel Bay that resembles the Gelidiums, Mr. Nott noticed it when he 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 not dichotomous. 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 the Fauschia with broad frond? If I did please save it for me.

Your obliged Friend, J.M. Weeks

8 FREMONTIA VOLUME 32:1, JANUARY 2004 Receipt of a reprint of the paper tion of a yearbook, she named it in which Laminaria ephemera was “Postelsia.” One of her students, eventually published by Setchell was Robert F. Griggs, found an abun- acknowledged by Mrs. Weeks on dance of an annual species of Lami- March 26, 1901, but it apparently naria growing near the station, and wasn’t until some months later (let- unaware that it had already been ter of November 25, 1901) that she described and named by Setchell, discovered with chagrin that in the proceeded to describe it exhaustively same paper Setchell had given the in the 1906 edition of “Postelsia.” name Weeksia to a new genus of red He named it Renfrewia parvula. algae. Previously, when Setchell had Griggs was impressed by the dis- suggested describing a new species coid holdfast vis-a-vis the stout with an epithet commemorating its branched holdfast of perennial spe- discoverer, Mrs. Weeks protested: cies of Laminaria and thus estab- “I am quite in earnest in not wish- lished a new genus to accommo- ing my name attached to any plant” date the local species and a previ- (letter of September 22, 1896). Mrs. ously described Japanese species. Weeks summed up the feelings of The holdfast of Laminaria ephem- all amateur collectors: “I shall re- era, however, is not strictly discoid, joice when definition and classifica- often having rudimentary branches tion are definitely & clearly settled” (haptera). Renfrewia currently is not (letter of June 26, 1899). Inciden- recognized as a genus separate from Professor Josephine E. Tilden returning tally, any shortcomings in Mrs. Laminaria. Weeks’s writing should not be as- from the third Pan Pacific Science Although Laminaria ephemera is Congress held in Tokyo, 1926. Use of cribed to lack of education: she delicate and shy, seeking sheltered photograph courtesy of S. Ducker. graduated from Knox College in pockets in the lowest intertidal and Galesburg, Illinois, in 1854, an un- upper subtidal zones, while Postelsia usual achievement for a woman of palmaeformis is rugged and resilient, that period. Her own explanation REFERENCES defying the strongest of waves, they strikes close to the heart of all ma- share similar stories of duplicated rine phycologists: “You must ex- Areschoug, J.E. 1853.Virginia, ett “discovery.” Even those seaweeds cuse this if you can’t read it readily nytt algslägte. Öfversigt af that are less distinctive have stories as five days of rising about 3 A.M. Förhandlingar Kongl. Svenska to tell. Victorians, who were in- Vetenskaps-Akademien 10:145-147. & an ill turn make writing jerky trigued, even obsessed by the sea- Griggs, R.F. 1906. Renfrewia parvula, business” (letter of June 20, 1897). side, its salubrious climate, and the a new kelp from Vancouver Island. The story of Laminaria ephem- Postelsia 1906:245-274. era does not end here, but shifts to inspiring array of creatures, were moved to write poems, of which the Howard, M.M. 1846. Ocean flowers and British Columbia, where Josephine their teachings. Binns and Goodwin. following, of uncertain origin but E. Tilden, a professor in the botany Bath, England. department at the University of found in M.M. Howard, “Ocean Mondragon J. and J. Mondragon. Minnesota, had established a bio- flowers and their teachings” (1846), 2003. Seaweeds of the Pacific Coast. logical station on the southwest is most appropriate: Sea Challengers. Monterey, CA. coast of Vancouver Island with the Ruprecht, F.J. 1852. Neue oder financial help of one of her devoted Oh! call us not weeds, but flowers unvöllstandig bekannte Pflanzen aus dem nördlichen Theile des Stillen students, Carolyn Crosby, whose of the sea. Oceans. Mémoires de l’Académie family owned Crosby Mills (later For lovely, and gay, and bright- Impériale des Sciences de Saint- General Mills). The station was near tinted are we! Petérsbourg, Sixième Série, Sci- Port Renfrew, a port used for ship- ences Naturelles 7(Bot.):55-82. ping logs from the adjacent forest. Our blush is as deep as the rose Setchell, W.A. 1901. Notes on algae. It was an idyllic site, with massive of thy bowers— I. Zoe 5:121-129. granite boulders covered with wave- Then call us not weeds, we are beaten seaweed, including the sea Ocean’s gay flowers. Paul C. Silva, University Herbarium, palm (Postelsia). This seaweed was UC, 1001 Valley Life Sciences Bldg. No. particularly striking, so that when 2465, Berkeley, CA 94720-2465. Tilden arranged for the publica- [email protected]

VOLUME 32:1, JANUARY 2004 FREMONTIA 9 Caulerpa taxifolia, a beautiful, aggressive invasive tropical green alga, appeared in a harbor near San Diego in 2000. Image courtesy of the University Herbarium, UC Berkeley.

CALIFORNIA’S NON-NATIVE SEAWEEDS by Kathy Ann Miller

he seaweed flora of Califor- flora, Marine Algae of California vide food, shelter, or companion- T nia is a world-class treasure. (Abbott and Hollenberg 1976) lists ship, or inadvertently, as hitchhik- The magnificent diversity and 669 species of red, brown, and green ers. For hundreds of years, our ter- abundance of seaweed populations seaweeds. The actual number is cer- restrial California has been gilded— reflect the dramatic sweep of our tainly much greater with the addi- and altered forever—by the unin- state’s rich coastal environments and tion of recently described species, tentional introduction of European habitats—from the Pacific North- species yet to be described, and spe- grasses; the list of introduced Medi- west to the subtropics, including cies that are currently arriving. Of terranean, South African, and Asian rocky shores and reefs, sandy those more recent introductions, weeds is enormously long. Pampas beaches, and offshore islands. Our some come to California naturally, grass! Yellow star thistle! Scotch scientific knowledge of California through range extensions due to El broom! Fennel! seaweeds is young, beginning with Niño events and long-term climate In the last few decades, we have the 1791 Malaspina expedition of dis- change, but some arrive via humans come to realize that our estuaries covery, growing through the efforts who unintentionally introduce non- and coastal oceans are also vulner- of European and east coast bota- indigenous species on hulls of ships able to introduced species, includ- nists, and coming of age with the or, for example, by improperly dis- ing invasive species that can cause collected works of W.A. Setchell and posing of aquarium plants. economic and ecological night- N. L. Gardner at the University of Non-native or exotic species, mares. There are an estimated 175 California at Berkeley. And yet, to especially those that spread like non-native species of plants and ani- this day, the delineation of species weeds, are both familiar and news- mals in San Francisco Bay alone, within our most common genera worthy. (Weeds, strictly defined, are and the number continues to grow. poses daunting challenges to phy- plants that live where they are not Rapacious invertebrates and fish cologists, who continue to pursue wanted and are ecologically or eco- lead the list and grab the news—the field, culture, and molecular studies nomically harmful.) Since the be- Asian clam, Chinese mitten crab, to unravel relationships at every taxo- ginning of civilization, humans have green crab, northern pike, striped nomic level. moved plants and animals to serve bass (an intentional introduction) The current comprehensive them, either intentionally to pro- are a few. But have you ever heard

10 FREMONTIA VOLUME 32:1, JANUARY 2004 of the infamous “Killer Weed,” distributions and it is hard to dis- as biological. Caulerpa taxifolia? There are inva- cern their history. These species are Finally, as every amateur and sive seaweeds here, too. known as “cryptogenic” because professional phycologist knows, Determining the presence of their original provenances are hid- many species look extremely simi- exotic seaweed species can be diffi- den from us. The ships that brought lar and are difficult to tell apart, cult if we have not witnessed the early explorers to our coasts were constituting complexes of what may introduction. We consider creatures wooden reefs supporting creatures be cryptic (hidden) or sibling to be native when they occupy the from every port visited over (closely related) species. Learning area in which they evolved; usually multiyear voyages, and were prob- more about our seaweed flora—and there are natural barriers to the dis- ably significant sources of introduc- detecting interlopers—is becoming persal of a species. Sometimes these tions, long before scientists estab- increasingly important. barriers shift through natural habi- lished a baseline for what is indig- The Mediterranean Sea, with tat or climate changes, allowing a enous. For example, Asparagopsis 85 introduced seaweeds, including species to expand its geographic taxiformis is a red alga, originally nine that are considered invasive, is range. Sometimes we are able to described from an Egyptian speci- a hot spot for introductions. At the access new areas via new technolo- men, but considered a cosmopoli- Thau Lagoon, an aquaculture site gies, such as scuba diving or tan member of subtropical and on the southern coast of France, 45 submersibles, and revise our records tropical communities worldwide. It species of seaweeds (23% of the for the ranges of species. But to occurs in southern California (San flora) are exotics, probably intro- distinguish introduced species from Diego, and the southern Channel duced with the Japanese oyster those that are native, we must rely Islands). Is it introduced? Has it Crassostraea gigas. In contrast, we’ve upon the history of collection data extended its range from Baja Cali- recorded a modest 12 non-native coupled with our understanding of fornia, Mexico? Or has it “always” seaweed species (or 2% of the sea- species distributions and natural dis- been here? We must critically re- weed flora) from our California persal of populations. visit nomenclatural history because coast. Most of these are diminutive When specimens from our coast the basis of our knowledge of sea- and easily overlooked, and several were sent back to European spe- weed distributions is cultural as well have been reported only from our cialists, vague collection notes (“in mare australe”) and mistakes as Seaweed scientific names, common names, simple as mislabeling were frequent, and authors and have left a legacy of confusion. For example, it was recently dis- or this article and those that covered and confirmed with mo- Ffollow, common names are lecular data that the type locality only provided if available. Unlike (original collection) of a very com- terrestrial plants, most seaweeds mon red seaweed known as Iridaea do not have colloquial names. For cordata was the tip of South America, purposes of precision, a scientific not Vancouver Island, Canada, the name may be followed by its provenance written on the her- author(s). If a species has been re- barium sheet. That species name is tained in its original genus, the therefore not applicable to our west author is the person who origi- coast species (now known as nally described the species, for ex- Mazzaella splendens, based on a Cali- ample, Postelsia palmaeformis fornia type specimen described by Ruprecht. If a species has been Setchell and Gardner). Similarly, transferred to another genus, the the southern California rockweed name of the describer will appear Label for only California collection Hesperophycus californicus was once in parentheses followed by the of Caulerpa taxifolia at the University Herbarium, UC Berkeley. Herbarium known as H. harveyanus, based on a name of the person who made the plant supposedly collected in labels rarely include the colloquial transfer. Thus, the citation name of the plant collected, but always Monterey in the 1830s—but prob- Caulerpa taxifolia (Vahl) C. Agardh include the scientific name with its ably collected in France and not indicates that this species was origi- author. Hesperophycus at all. nally described by Vahl (in 1802 as Furthermore, many of our com- Fucus taxifolius) and transferred to mon seaweeds have extraordinarily Caulerpa by C. Agardh (in 1817). broad “cosmopolitan” geographic

VOLUME 32:1, JANUARY 2004 FREMONTIA 11 own invasion hot spot, San Fran- ent reproductive characteristics, are Caulacanthus has been observed at cisco Bay. recorded from San Francisco Bay. numerous sites during the last de- Good examples are Aglaotham- One is entirely asexual, a good can- cade or so, but was not recorded nion tenuissima (Bonnemaison) didate for an introduced species during extensive Bureau of Land Feldmann-Mazoyer and Polysiphonia (no need to find a mate). And by Management surveys 30 years ago. denudata (Dillwyn) Greville ex the way: Bryopsis hypnoides Just last year, I spotted a large red Harvey, both small red filaments Lamouroux, reported from brown patch in the intertidal as I originating in England and detected Humboldt Co. to San Pedro, was walked my sister’s dog in a park on in San Francisco Bay. More re- originally described from the San Francisco Bay. This proved to cently, another small red filamen- Mediterranean. Like so many Cali- be the first sighting of this species tous species, Polysiphonia harveyi J. fornia cryptogenic species with far- at the Bay. Where did it come from? Bailey, once thought to be a native flung distributions, we have to How does it disperse? Are these of the east coast of North America, wonder: native or introduced? new introductions, or is this species has been traced through molecular The red alga Caulacanthus extending its range due to climate methods as originating in Japan and ustulatus (Mertens ex Turner) change? spreading, via several introduc- Kützing is a cosmopolitan, crypto- Lomentaria hakodatensis Yendo tions, to the Atlantic, including the genic species, reported from Japan, is an inconspicuous red alga that British Isles, New Zealand, and US, Indonesia, South Africa, Spain, New has quietly spread from its native including California, where it has Zealand—all places where plants Japan to the Mediterranean, Phil- masqueraded as Polysiphonia have been designated as types for ippines, Hawaiian Islands, Austra- acuminata Gardner and P. simplex species now considered synonyms lia and the west coast from British Hollenberg. A great example of a of C. ustulatus. This small, turfy spe- Columbia, Washington, and Or- sibling/cryptic species complex. cies also occurs in Australia, Africa, egon to California and Mexico. We Another example of a sibling India, and Hawaii, as well as British assume it is an exotic because it species complex that is yet to be Columbia, Washington, and Cali- wasn’t reported in early surveys teased apart is the green algal ge- fornia. (As a student in 1979, I re- (e.g., E. Yale Dawson’s surveys of nus Bryopsis. In addition to Bryopsis member being shown a small popu- the Northern coast from Cape corticulans Setchell, three lation on Vancouver Island. Exotic! Mendocino to Crescent City). Be- bryopsidoid entities, with differ- Exciting!) In southern California, cause many of our California spe-

Illustrations from the DeCew Guide

hroughout this article and within other articles in this Fremontia T issue, illustrations appear from Tom DeCew’s Guide to the Seaweeds of British Columbia, Washington, Oregon, and Northern California, a Web version of a book on northeast Pacific seaweeds conceived by the late Tom DeCew, and brought to its present state of completion with the collaboration of Paul Silva, Richard Moe, and Robert Rasmussen. DeCew planned the book as a detailed complement to Marine Algae of California by I.A. Abbott and G.J. Hollenberg (Stanford University Press, 1976). For each of the species of macroalga (seaweed) that occurs relatively commonly in British Columbia, Washington, Oregon, or northern California, Decew assembled information from specimens in West Coast herbaria and from personal collections. Each page contains a graphic depiction of distribution and reproductive status, often along with a specially commissioned original line drawing. He also compiled all published references of scientific studies for each species. The Web version, which follows the layout of the book, including all of the data and 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.

12 FREMONTIA VOLUME 32:1, JANUARY 2004 cies also occur in Japan, it is diffi- cult to determine without good his- torical collections if a given species occurs naturally throughout the eastern and western north Pacific or if it is an introduction. Gelidium vagum Okamura is another small red alga, native to Japan, China, and Russia. It has a limited distri- bution in British Columbia and has now appeared in Tomales Bay, Cali- fornia. Since Tomales Bay is home to oyster mariculture, this species may have been introduced as a hitchhiker with oyster spat from Washington. As in the Thau Lagoon, oysters have been the vector for many an introduced species, including Sar- gassum muticum (Yendo) Fensholt, a species that can be considered our first truly invasive species due to its The distinctive “Dead man’s fingers,” Codium fragile subsp. tomentosoides, has the rapid spread. Introduced to the west second, well-earned, name of “oyster thief.” Our native subspecies lives on the outer coast from Japan before World War coast, while the invasive C. fragile subsp. tomentosoides has been found only in San II, it was detected in British Co- Francisco and Tomales bays. Line drawing from DeCew Guide, used with permission lumbia in 1944, Oregon in 1947, from P. Silva. Crescent City, California in 1963, Santa Catalina Island in southern tic until a weedy strain (subsp. seaweeds are infamous aggressors, California in 1970, and San Fran- tomentosoides) was introduced to the though how they will play in Cali- cisco Bay in 1973. While studies Netherlands from Japan (its pre- fornia is yet to be seen. In 2000, have not confirmed that Sargassum sumed source) shortly before 1900. both the green alga Caulerpa taxifolia muticum is ecologically harmful, This weed has spread with increas- (M. Vahl) C. Agardh, and the brown (e.g., displacing native species), it is ing rapidity throughout the North kelp Undaria pinnatifida (Harvey) certainly a conspicuous and abun- Atlantic, including the Mediterra- Suringar showed up in southern dant part of our intertidal and shal- nean and the North American coast California. News of the first has low subtidal communities. In 1973, from Nova Scotia to North Caro- swamped interest in the second, when it arrived in the Solent region lina. It is now at home in New because Caulerpa had proven of southern England (possibly from Zealand, Australia, and California itself to be a devastating plague in a shipment of British Columbia oys- (San Francisco and Tomales bays). the Mediterranean. Distasteful to ters), efforts to eradicate it ended in An herbivorous sea slug checks the herbivores, it has covered acres of failure, even after volunteers re- growth of the weed in New Zealand. seafloor, out-competing native moved 475 tons over the course of In England and Ireland, the weed seagrasses and smothering native three years. Such efforts here, if we has been reported to displace a na- fauna. chose to undertake them 30 years tive species, Codium tomentosum. In The culprit is an escaped inva- after the introduction, would surely New England it overgrows and dis- sive aquarium strain, identified as fail as well. lodges shellfish (thus earning the originating in Australia and subse- Oyster mariculture may be im- name “oyster thief”) and gangs up quently hybridizing with other plicated in the dispersal of the green with an introduced bryozoan to dis- strains, creating a strain tolerant of alga Codium fragile subsp. place native kelps. To date in Cali- temperate waters. During eelgrass tomentosoides (van Goor) P.C. Silva, fornia it is confined to two bays and surveys, Caulerpa was discovered in another weed adept at asexual re- is thus separated from our native a shallow lagoon, Agua Hedionda, production. Codium fragile com- subspecies, which is well repre- near San Diego and in Huntington prises several well-behaved subspe- sented on the outer coast from Harbor, California. Because the cies native to temperate Pacific Alaska to Baja California, Mexico. plant fragments and regenerates, it coasts, but was lacking in the Atlan- The next two non-native cannot be manually removed.

VOLUME 32:1, JANUARY 2004 FREMONTIA 13 real until all species of Caulerpa are Undaria pinnatifida, native to banned from the aquarium trade. Japan, Korea, and China, is known While news of Caulerpa reached as wakame in Japanese cooking. the public in newspapers, television, Photograph by K.A. Miller. and Internet reports, Undaria pinnatifida, a two meter long kelp native to Japan, Korea, and China silently skipped its way up the Cali- fornia coast from harbor to harbor, from Los Angeles to Santa Barbara. Protect Our Coast from An In 2001, it leaped to Monterey Har- Invasive Kelp bor and a small cove on the leeside of Santa Catalina Island. Over the ndaria pinnatifida originated in Asia and has spread past several decades, Undaria has in- Uaround the world to Australia, New Zealand, South America, vaded Europe, the Mediterranean, Europe—and now California. Undaria is unlike any other California South America, and Australia and kelp. Its blade is delicate (thin), elaborately lobed along the margin, New Zealand, through deliberate and has a prominent midrib. It can be three to five feet long, and if you introductions (it is a prized food) look closely, you’ll see tiny dots scattered over the surface, formed by and probably by hitchhiking on the tufts of microscopic hairs. The reproductive structure (sporophyll) hulls of ships. Like many invaders, it develops below the blade, just has a broad tolerance for tempera- above the holdfast, and is unique ture and thrives in quiet water, espe- in that it is deeply frilled, like old- cially in the presence of the nitrogen fashioned ribbon candy. The blade enrichment typical of harbors. In persists from spring through sum- California harbors, it is a short-lived mer, and then erodes back to the annual, recruiting in the early spring, holdfast and sporophyll. and with a smaller pulse of new If you find this species, take a sporophytes in the fall at some sites, picture of it underwater. If this is and then disappearing for the win- not possible, collect a specimen ter. Santa Catalina Island, one of the for a photograph but do not throw California Channel Islands, is the Holdfast of Undaria pinnatifida. it back into the water! Rather, dis- only site at which it co-occurs with pose of it on land, and contact Dr. native seaweeds in a relatively natu- Kathy Ann Miller, Wrigley Ma- ral (non-harbor) setting. There it rine Science Center, Santa lives at record-setting depths (up to Catalina Island, (310) 510-4012 24 m) in the clear offshore waters. In ([email protected]). three years of observation at Catalina, There are several websites with Undaria has spread within the site, useful additional information moving into shallower depths (in- about this seaweed (search the cluding the understory of a giant Web for Undaria pinnatifida). For kelp bed), but it has not extended its example, the site at biology.ucsc.edu/ range to other sites around the is- people/potts/lonhart/undaria.html land or other islands. includes many images of this plant. Blade of Undaria pinnatifida. Because it is an annual, Undaria does not appear to compete with perennial native kelps, but it cer- Caulerpa patches were covered with eradication can be declared. tainly warrants status as a potential tarp and bleach was injected beneath. Ten species and four subspecies pest, capable of usurping habitat for A couple of years of this treatment of Caulerpa are sold in aquarium much of the summer. The Califor- (and several million dollars) resulted stores in southern California. Of nia Department of Fish and Game in a tentative victory: Caulerpa has these, 12 are capable of surviving in is making efforts to eradicate it from not been seen at these sites in more the temperate zone, and are poten- Santa Barbara harbor, and volun- than a year, and has not been re- tial escapees.Because of difficulties teer groups are chipping away at ported from new sites. Quarterly identifying the invasive strain, the the Catalina and Monterey popula- monitoring continues until complete chance of a new introduction is very tions. However, many predict that

14 FREMONTIA VOLUME 32:1, JANUARY 2004 any effort to remove Undaria at this that might indicate that it has settled introduced macrophytes. Marine point is too little and too late. The in. This too was floating, like the Pollution Bulletin 44:32-38. kelp life history promotes long-term ecad (ecological form) of Ascophyllum Carleton, J.T. 2001. Introduced species survival of the species through called scorpioides; attached plants have in US coastal waters: environmental highly variable conditions. Each not yet been observed. impacts and management priorites. Pew Ocean Commission, Arling- kelp plant is capable of dispersing This example reminds us that ton, Virginia. www.pewoceans.org millions of spores and establishing the establishment of a non-native DeCew, T. (see sidebar on page 12) populations of the microscopic, fila- species is not a simple slam dunk. Guide to the Seaweeds of British mentous sexual phase. Undaria is Those species blessed with a pecu- Columbia, Washington, Oregon, probably here to stay, although we liar suite of attributes (broad physi- and Northern California. Posted should make every effort to eradi- ological tolerances, copious repro- to the UC/JEPS website at cate it. Whether it will continue its ductive abilities, and facile dispersal ucjeps.berkeley.edu/guide/ march up the coast remains to be mechanisms) still require a large dose Deysher, L. and T. A. Norton. 1982. seen. of luck to establish on foreign shores. Dispersal and colonization in Sar- gassum muticum (Yendo) Fensholt. A final species deserves men- To date Sargassum muticum, tion, because there is evidence that Journal of Experimental Marine Bi- Caulerpa taxifolia, Undaria pinnatifida, ology and Ecology 56:179-195. it may be establishing itself in Cali- and perhaps Caulacanthus ustulatus McIvor, Lynne, Christine A. Maggs, fornia after years of unsuccessful can be considered invasive weeds as Jim Provan and Michael J. introductions. This is the brown well as non-native components of Stanhope. 2001. rbcL sequences re- alga Ascophyllum nodosum (Linnaeus) the California flora. What will the veal multiple cryptic introductions Le Jolis, the knotted wrack native future hold? Introductions seem to of the Japanese red alga Polysiphonia to the Atlantic, used to steam clams be increasing (Caulerpa and Undaria harveyi. Molecular Ecology 10:911- and lobsters or pack bait. It has been arrived simultaneously in the new 919. sighted floating in the Bay on many millennium), and the consequences Silva, P.C. 1979. The benthic algal flora of central San Francisco Bay. occasions over the years, surely as of this global homogenization of spe- Pp. 287-345 in San Francisco Bay: discarded packing material, but only cies are mixed. Some, or perhaps this year has a sample been collected The Urbanized Estuary, T. J. most, introductions are benign or Conomos, editor. Pacific Division, neutral, but others are disastrous, American Association for the The brown alga Ascophyllum nodosum, setting the scene for “invasional native to the Atlantic, is used to pack bait. Advancement of Science. San Fran- It may be establishing itself in California meltdowns,” facilitating other inva- cisco, CA. after many years of unsuccessful sions and sometimes increasing the Silva, P. C. 1996. California seaweeds introductions. Image courtesy of impact of subsequent invaders. I’ve collected by the Malaspina Expe- University Herbarium, UC Berkeley. often thought that the urchin-domi- dition, especially Pelvetia (Fucales, nated habitats that have replaced Phaeophyceae). Madroño 43:345- 354. some of our kelp forests in southern Silva, P. C., R. A. Woodfield, A. N. California are fine settings for the Cohen, L. H. Harris, & J. H.R. invasion of seaweed species distaste- Goddard. 2002. First report of the ful to urchins, who could really chal- Asian kelp Undaria pinnatifida in lenge our native kelps to a run for the northeastern Pacific Ocean. their money. But luck has been with Biological Invasions 4:333-338. us, so far. It’s clearly more impor- Verlaque, M. 2001. Checklist of the tant than ever to continue to study macroalgae of Thau Lagoon our shores and to train our students (Hérault, France), a hot spot of to recognize our native California marine species introduction in Europe. Oceanologica Acta. 24:29-49. seaweeds—and the new arrivals. Williams, S. L., and E. D. Grosholz. 2002. Preliminary reports from the Caulerpa taxifolia invasion in south- REFERENCES ern California. Marine Ecology Progress Series 233:307-310. Abbott, I.A. and J. Hollenberg. 1976. Marine Algae of California. Stanford Kathy Ann Miller, Wrigley Marine Sci- University Press. 827 pp. ence Center on Santa Catalina Island, Boudouresque, C. F. and M. Verlaque. University of Southern California. 1 Big 2002. Biological pollution in the Fisherman Cove, Avalon, CA 90704 Mediterranean Sea: Invasive versus [email protected]

VOLUME 32:1, JANUARY 2004 FREMONTIA 15 CALIFORNIA SEAWEED INDUSTRY by Paul C. Silva

eaweed industry in Califor- tition from other sources of potash Gelidium, a red alga. The first agar S nia was born of necessity. caused the plant to close soon after factory in California was established Prior to the First World the war ended. in 1920 in Glendale, but operations War, the mines of Stassfurt, Ger- The Second World War also were soon moved to San Diego. many, were essentially the world’s produced a crisis by cutting off our Initially, the chief function of the sole source of potash, which is used supply of agar, normally imported industry was to purify Japanese agar in making fertilizers, fireworks, and from Japan. Agar is one of three to bring it up to bacteriological stan- explosives. To replace this source commercially important water- dards, but with the supply cut off by in wartime, the Hercules Powder soluble polysaccharides found in the war, primary production was Company established a plant on the seaweed. Although it is principally essential. The source was Gelidium shore of San Diego Bay to obtain known as the gelling agent of the robustum, which could be obtained potash (and other chemicals) by in- substrate on which bacteria are in large quantities from fishermen cinerating giant kelp (Macrocystis grown, it also has many industrial in northern Baja California, where sp.). At the peak of activity, there uses. Prior to 1940, it was extracted it grows abundantly in the lowest were 1,100 employees, but compe- mainly from various species of intertidal and shallow subtidal zones. Agar production in San Diego ceased about 1986 with the closing of the American Agar and Chemical Company, but the indus- try continues in Ensenada, Baja California. The other two water-soluble polysaccharides (“phycocolloids”) are alginate (the sodium salt of alg- inic acid), obtained from large brown seaweeds (kelps and rockweeds), and carrageenan, obtained from certain red seaweeds. All three phycocolloids are emulsifiers and are often used interchangeably, but they differ in certain physical properties so that each 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 as textile sizing and ink manufacture. An alginate industry was started in San Diego in 1927 and thrives today under the name ISP (International Specialty Products) San Diego. The former Kelco Company figures prominently in the local history of this industry. The beds of Macrocystis that extend from San Diego to Monterey provide the raw material. This Kelco ship harvests and processes the tops of the giant kelp, Macrocystis A small factory for processing pyrifera. The colloidal extract, sodium alginate, is used as a thickener in paints, carrageenan was operating in the dyes, beer, and canned foods. Use of photograph courtesy of J. Hansen. San Francisco Bay area during the

16 FREMONTIA VOLUME 32:1, JANUARY 2004 Along with commercial, large scale harvesting, Porphyra sp., known as nori, is collected in northern California as part of a cottage industry. 1930s, using Mazzaella spp. as a source, but the world market is now abundantly supplied, often satu- rated, by the output of seaweed farms in many tropical countries, especially the Philippines, where carrageenan-rich Eucheuma spp. is grown in ponds or on reefs close to shore. Historically, carrageenan was extracted from Chondrus crispus (“Irish Moss”) by coastal peoples in the North Atlantic and used to thicken puddings. In Japan, commercial produc- tion of foodstuffs from seaweed, es- pecially amakusa nori from Porphyra, kombu from Laminaria, and wakame from Undaria, is big business, but no such industry has been established in California.

Paul C. Silva, University Herbarium, UC, 1001 Valley Life Sciences Bldg. No. 2465, Berkeley, CA 94720-2465. A less utilitarian use of seaweed: toys made from bull kelp (Nereocystis luetkeana), with [email protected] the air bladders forming perfect doll heads. Use of photograph courtesy of K. Hubbard.

VOLUME 32:1, JANUARY 2004 FREMONTIA 17 Corallina vancouveriensis, a coralline alga, is a common inhabitant of the mid to low intertidal. Coralline algae are impregnated with calcium and may form a crust, be erect with jointed fronds, or both. Photograph by S. Sharnoff.

DISTRIBUTION OF CALIFORNIA SEAWEEDS by Paul C. Silva

enthic marine algae, or the continent, for the most part hid- to pop the floats of Macrocystis and B “seaweeds” as those of us den from ordinary view. Few people use the Egregia as a jump rope. Thus who know them on a first- have experienced the glorious sight the seaweed experience of most name basis prefer to call them, are of a rocky shore covered with a luxu- Californians would barely hint at those algae that are attached to riant growth of seaweeds exposed the glorious diversity and beauty to rocks, other algae, animals, or man- at extreme low tide. In California, be found in this fascinating group made structures such as wood pil- such a sight is possible only very of plants. ings. They are big enough to see early on summer mornings or just and feel, although it is necessary to before dark in winter. These times use a hand lens or microscope to are dictated by the annual tidal cycle VERTICAL determine the essential structure of and are not likely to be chosen by DISTRIBUTION many small species. While their eco- the casual visitor to the shore. In nomic importance is insignificant addition, most people head for Seaweeds do not venture up- compared to the role played by the sandy beaches, where they encoun- ward beyond the splash zone be- microscopic phytoplankton at the ter rotting masses of seaweed cause they must be washed by the base of the great ocean food web, washed ashore, often giant kelp sea at least once each day. Few spe- their beauty and diversity have (Macrocystis) intertwined with cies can tolerate the long periods aroused the passion of naturalists feather boa kelp (Egregia), teeming of exposure imposed at high tide for the past two centuries. with hungry critters, such as kelp levels. As the tide level decreases, Seaweeds cling to the edge of flies and sand hoppers. Children like the number of species increases

18 FREMONTIA VOLUME 32:1, JANUARY 2004 markedly, the greatest diversity be- brown, and green), each of which FLORISTIC ANALYSIS ing found at the lowest tide level has a characteristic complement of (about minus 0.6 meter on outer photosynthetic pigments that is ef- The coast of California is home coasts of California) and in the ad- fective in normal light, are adapted to about 670 species of marine al- jacent subtidal fringe. Some species to living in deep water by the pos- gae, a mere 12% of the number of live only in the subtidal, at depths session of accessory pigments with higher plant species. Endemics limited by the availability of light. appropriate absorption spectra. (those species restricted to Califor- When the water is exceptionally At very low tide, the exposed nia) account for 14% compared to clear, a few species may grow at shore often appears to be divided 24% for higher plants (Hickman depths as great as 268 meters, as in into several horizontal zones, dif- 1993). By far the largest group of the Bahamas (Littler et al. 1986). In fering in color and texture depend- species (44%) are those that are es- California, the Channel Islands are ing on the species that find optimal sentially restricted to the Pacific noted for the clarity of the sea, be- growth conditions at those particu- Coast of North America, with ing correlated with the absence of lar tide levels. Generally, one of the ranges extending from some point river runoff. higher zones is comprised of mem- north of California (some as far With increasing depth of water, bers of the Fucus family, which are north as the Bering Sea) southward. the intensity of light decreases commonly called “rockweeds,” This group includes mostly cold- sharply while the quality of light while the lowermost zone is domi- temperate species, but also some changes dramatically. Red light is nated by members of the order that survive or even thrive in mod- completely absorbed in the upper Laminariales, commonly called erately warm waters. Another group few meters so that a blue-green twi- “kelps.” The subtidal fringe is often (21%) are those that are essentially light prevails in deeper water. The bright pink from encrusting calcar- restricted to southern California and various classes of seaweeds (red, eous algae of the order Corallinales. adjacent Baja California, constitut- ing a warm-temperate flora. An equal number of species (21%) were originally described from areas other than the North American Pacific Coast and include so-called cosmopolitan species, introduced invasive species (“weeds” in the usual sense of the word), and many species with notably disjunct distri- butions that need to be confirmed by close taxonomic scrutiny. Many genera and even some species of the California marine flora are be- lieved to occur also in Japan, but future molecular studies will prob- ably reveal at least slight genetic differences between western and eastern Pacific populations.

TEMPERATURE: PRIMARY FACTOR IN DETERMINING DISTRIBUTION

Temperature is the overriding factor in determining the geo- graphic distribution of seaweeds, while oceanic currents impose and Fucus gardneri, a rockweed, is a common inhabitant of the high intertidal zone. Line maintain the temperature regimes. drawing from DeCew Guide, used with permission from P. Silva. The effect of the Gulf Stream on

VOLUME 32:1, JANUARY 2004 FREMONTIA 19 Oceanic currents in the Atlantic Ocean. Oceanic currents in the Pacific Ocean. Note particularly the Kuroshio Current system, Note particularly the Gulf Stream, which which brings warm water from the equator to the coast of Japan but ends as the brings warm water from the equator to relatively cold California Current. the eastern coast of North America and the Atlantic coast of Europe. climate is common knowledge. It The temperature of the cur- seasons, but is pushed downward originates as the North Equatorial rents is sharply reduced under con- by upwelled water and moves up- Current and passes through the ditions favorable to upwelling. In ward when upwelling ceases. Straits of Florida with sufficient this process, prevailing westerly In the southern hemisphere, the velocity to carry the warm water winds with cliffs to the left (in the Coriolis force deflects the move- along the Atlantic coast of the northern hemisphere) push water, ment of water to the left, away United States as far as Cape Cod which has been warmed by the sun, from the coast as in the northern and across the ocean to the British along the coast. This forward hemisphere since the cliffs that Isles and Norway. Equally spec- movement is subjected to the Co- confine the winds are on the right. tacular, but in reverse, is the Peru- riolis force (an effect of the rota- Upwelling along the coasts of Chile vian Current (formerly called the tion of the earth), which causes the and Peru lowers the temperature Humboldt Current), by which very warm water to be deflected to the of the Peruvian Current while up- cold water moving clockwise in the right, away from the coast. Cold welling in Namibia and Atlantic Southern Ocean is deflected by the water then wells up from moder- South Africa lowers the tempera- tip of South America, sending it ate depths. Low-lying coasts allow ture of the Benguela Current. On northward along the Pacific coast the winds to spread out and dimin- the west coast of both South and imposing temperate conditions ish in strength, while headlands America and Africa, kelps, which at tropical latitudes. Of greatest confine the winds and thus inten- are quintessentially cold-water importance in the North Pacific is sify the effect of upwelling. In plants, grow within the tropics. the Kuroshio Current system. Like northern California and northern Whether a current is warm or the Gulf Stream, it originates as Baja California, intense upwelling cold is relative. While the Alaska the North Equatorial Current and begins in March and continues Current is sufficiently warm to proceeds clockwise past the Phil- through July, with water tempera- ameliorate the climate of southern ippines and Taiwan along the coast tures in the spring lower than those Alaska, the California current, en- of southern Japan, where it is called of winter. When the prevailing forced by upwelling, is cold in its the Kuroshio Current. North of north-northwest winds diminish northern reaches. Inshore water Japan, it merges with the easterly and upwelling ceases in autumn, a temperatures reach their maximum drift of the North Pacific Ocean. relatively warm surface counter- in late August and early Septem- On reaching North America, part current (the Davidson Current) ber, but only in southern Califor- of the current is deflected north- flows northward along the coast, nia do they reach 20ºC or slightly ward in a counterclockwise gyre in being especially noticeable from higher. In northern California the the Gulf of Alaska (Alaska Cur- November through January. This usual annual range is 9-16ºC. rent) while part is deflected south- counter-current is generated by the Whereas the distribution of ward as the California Current. California Current and flows in all those species that grow in the in-

20 FREMONTIA VOLUME 32:1, JANUARY 2004 tertidal and shallow subtidal zones biologists to be algae, and by other is controlled by the temperature of biologists to be bacteria). the surface water, those that live in deeper waters usually have a par- ticular depth range correlated with GEOGRAPHIC temperature. For example, the kelp DISTRIBUTION Agarum fimbriatum grows in the shallow subtidal zone in the North- The biogeography of the ma- west, but it occurs in California rine algae of California is a topic only in deep water (to about 115 that has engaged me for more than meters) in the Channel Islands, 50 years. At the time that I began being inhibited by the warm sur- graduate work (1946), there was a face water. widely held belief that Point Con- ception caused a major break in the marine biota, but there was no SECONDARY published evidence concerning FACTORS IN seaweeds. The idea, which is largely DETERMINING zoological, is based on the lifelong DISTRIBUTION work of William Healey Dall (1845-1927), a malacologist (Dall Oceanic currents along the west coast of 1916). I doubt that during Dall’s North America. Note that the California When a species is within its lifetime any significant collecting Current flows southward past Pt. acceptable temperature range, its Conception into northern Mexico before of molluscs or any other form of distribution may be limited by lack reversing directions and sending a branch marine life was done in the vicinity of a suitable habitat. Factors influ- northward into the Southern California of Point Conception. The coast- Bight. encing suitability include availabil- line from Gaviota (just west of ity of a firm substrate, exposure to Santa Barbara) northward to neither in Goode’s California Place wave action, exposure to shifting Carmel Bay was nearly inaccessible Names nor in Jepson’s index of geo- sand and gravel, and salinity. Some prior to 1940. Highway 1 was built graphical names. Reed extended species (called cumaphytes, or surf between Carmel Bay and San Luis the range of several species hith- plants) are restricted to the most Obispo County in the late 1930s, erto known only as far south as heavily exposed sites. Others pre- but was closed during the war. Monterey, and this information fer semi-protected reefs covered by When I became immersed in the would have been useful to future sand. Most species have a particu- study of marine algae, I soon found workers had it been published. In lar niche, such as surf grass beds at that the only significant collections the absence of published informa- zero tide-level, or the vertical sides from along this stretch of coast tion on this stretch of coast, gener- of boulders. Most species can sur- were those made by (and for) alizations on the role of Point Con- vive short periods of lowered sa- Rhoda Ramona Reed, a master’s ception in biogeography were in linity, but few can tolerate consis- student of Professor William fact based on comparisons between tently brackish water. For example, Albert Setchell at Berkeley. Reed’s the biota of Santa Barbara and that many conspicuous species com- family owned a ranch near Lion of the Monterey Peninsula. monly found on the outer coast of Rock on the northwest shoulder of During the period 1946-1952, central California are missing in Point Buchon in San Luis Obispo at the suggestion of Gilbert Mor- San Francisco Bay, and those that County. Collections from there gan Smith, my major professor at are there penetrate to a distance and from “Oilport” were reported Stanford University, I collected in- from the Golden Gate in direct in a thesis that was submitted in tensively along the California coast, proportion to their tolerance of May 1917 but was never published. as well as on the Channel Islands brackish water (Silva 1979). Wher- Reed gave the location of her of southern California and the ever a creek discharges fresh water father’s ranch as “Pecho,” and in- mainland and coastal islands of into the intertidal, the discharge deed there is a Pecho Creek and northern Baja California. It turned will be accompanied by a brackish- Pecho Rock nearby. “Oilport” was out that Point Conception does tolerant flora composed of bright a facility built in 1907 east of Port indeed play an important role in green strands (Enteromorpha) and San Luis in San Luis Obispo Bay the geographic distribution of Pa- blackish green carpets (blue-green at a site now occupied by Shell cific coast marine algae (and assur- prokaryotes, considered by some Beach. Inexplicably, “Oilport” is edly marine animals as well), but

VOLUME 32:1, JANUARY 2004 FREMONTIA 21 Table 1. Southern end–points of conspicuous cold–temperate seaweeds restricted to the mainland of California (but including the Farallon Islands). Counties arranged from north to south.

Humboldt County Farallon Islands unspecified Mazzaella parksii (Setchell & Gardner) Acrosiphonia spinescens (Kützing) Kjellman Hughey et al. Porphyra papenfussii Krishnamurthy Ulva conglobata Kjellman Porphyra pseudolanceolata Krishnamurthy Trinidad Santa Cruz County Palmaria hecatensis Hawkes Greyhound Rock Cape Mendocino Plocamium oregonum Doty Sphacelaria racemosa Greville Monterey County Mendocino County Monterey Peninsula Point Cabrillo (near Caspar) Arthrocardia silvae Johansen Coilodesme bulligera Strømfelt Fauchea fryeana Setchell Weeksia coccinea (Harvey) Lindstrom Hollenbergia subulata (Harvey) Wollaston Monostroma grevillei (Thuret) Wittrock Sonoma County Porphyra abbottiae Krishnamurthy Shell Beach Porphyra miniata (C. Agardh) C. Agardh Porphyra torta Krishnamurthy Porphyra schizophylla Hollenberg Bodega Head and Bay Pylaiella tenella Setchell & Gardner Cumathamnion sympodiophyllum Wynne & Rosenvingiella constricta (Setchell & Gardner) Daniels Silva Odonthalia washingtoniensis Kylin Serraticardia macmillanii (Yendo) Silva Porphyra sanjuanensis Krishnamurthy Stictyosiphon tortilis (Ruprecht) Reinke Point Lobos Marin County Chiharaea bodegensis Johansen Tomales Bay Malpaso Creek Chaetomorpha cannabina (Areschoug) Porphyra smithii Hollenberg & Abbott Kjellman Kasler Point Fucus spiralis Linnaeus Lessoniopsis littoralis (Tilden) Reinke Duxbury Reef Porphyra pulchra Hollenberg Laminaria groenlandica Rosenvinge Jade Cove Odonthalia lyallii (Harvey) J. Agardh Laminaria ephemera Setchell Porphyra fucicola Krishnamurthy Little Sur River Ulvaria obscura var. blyttii (Areschoug) Bliding Isabbottia ovalifolia (Kylin) Balakrishnan Point Sur San Francisco County Pleurophycus gardneri Setchell & Saunders San Francisco (City) and San Francisco Bay Big Sur River Acrosiphonia saxatilis (Ruprecht) Vinogradova Ulva stenophylla Setchell & Gardner Elachista fucicola (Velley) Areschoug Lola lubrica (Setchell & Gardner) A. Hamel San Luis Obispo County & G. Hamel unspecified Percursaria percursa (C. Agardh) Acrosiphonia coalita (Ruprecht) Scagel Rosenvinge et al. Porphyra kanakaensis Mumford Porphyra lanceolata (Setchell & Hus) G.M. Ralfsia fungiformia (Gunnerus) Setchell & Smith Gardner San Simeon Urospora doliifera (Setchell & Gardner) Doty Hedophyllum sessile Setchell

22 FREMONTIA VOLUME 32:1, JANUARY 2004 Table 1. (cont.)

Cayucos Santa Barbara County Porphyropsis coccinea (Areschoug) Rosenvinge Point Sal Atascadero State Beach Dictyoneurum californicum Ruprecht Bornetia californica Abbott Erythrophyllum delesserioides J. Agardh Montana de Oro State Park Point Conception Hymenena multiloba (J. Agardh) Kylin Ahnfeltiopsis linearis (C. Agardh) Silva & Kornmannia leptoderma (Kjellman) Bliding DeCew Macrocystis integrifolia Bory Analipus japonicus (Harvey) Wynne Percursaria dawsonii Hollenberg & Abbott Bossiella plumosa (Manza) Silva Pylaiella gardneri Collins Constantinea simplex Setchell Lion Rock Dilsea californica (J. Agardh) Kuntze Pelvetiopsis limitata (Setchell) Gardner Fucus gardneri Silva Postelsia palmaeformis Ruprecht Halosaccion glandiforme (S.G. Gmelin) Diablo Cove Ruprecht Alaria marginata Postels & Ruprecht Hymenena flabelligera (J. Agardh) Kylin Callophyllis crenulata Setchell Mastocarpus jardinii (J. Agardh) J. West Clathromorphum parcum (Setchell & Foslie) Odonthalia floccosa (Esper) Falkenberg Adey Phaeostrophion irregulare Setchell & Nereocystis luetkeana (Mertens f.) Postels & Gardner Ruprecht Gaviota Shell Beach Gloiosiphonia verticillaris Farlow Hymenena fryeana Kylin Goleta Pismo Beach Palmaria mollis (Setchell & Gardner) van Farlowia conferta (Setchell) Abbott der Meer & Bird

not in the simplistic way the story was usually told. The kelp Dictyoneurum califor- At Point Conception, the coast nicum is common between Cape veers sharply eastward toward Mendocino and Monterey, but is found as far north as British Santa Barbara. The relatively cold Columbia and as far south as Santa California Current continues its Barbara County. Line drawing southward sweep, bathing the from DeCew Guide, used with northern Channel Islands and then permission from P. Silva. flowing into Mexican latitudes where it is warmed. Part of it then returns northward, bathing the southern Channel Islands with rela- tively warm water and continuing counter-clockwise in the Southern California Bight. The other part continues southward as a weak cur- rent. The relatively warm water in the Bight creates a sharp tempera- ture gradient immediately east of Point Conception so that the Santa Barbara flora is indeed very different from that of the Monterey Peninsula. The ranges of cold-temperate

VOLUME 32:1, JANUARY 2004 FREMONTIA 23 species are related to temperature rather 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 south as Lion Rock in San Luis Obispo County. Some species drop out at Point Conception or intrude a short distance into the Bight. Many spe- cies, however, continue their distri- bution on the northern Channel Is- lands (see Table 2). Some of these species skip the southern Channel Islands but appear again on the mainland of northern Baja Califor- nia in pockets of upwelled cold wa- ter (see Table 3). Laminaria setchellii, Hedophyllum sessile, a distinctive kelp, is found from Alaska to Point Conception, but for example, occurs on the Califor- has yet to be found between Santa Cruz and Point Sur. Line drawing from DeCew nia mainland as far south as Point Guide, used with permission from P. Silva. Conception, then on the northern Channel Islands, with the south- flora is compared with a warm- including a remarkable number of ernmost locality being Punta Banda, water flora. Throughout the world, members of the order Rhody- just south of Ensenada, Mexico. large fleshy brown and red algae are meniales, are totally subtidal accord- Some species skip the Channel Is- generally restricted to cold water. ing to data given by Abbott and lands as well as southern Califor- An excellent overview of the Hollenberg (1976). Their vertical nia, reappearing in upwelling areas seaweeds of southern California is ranges are variable. A few species of northern Baja California (see given by Murray and Blair (1993), occur in the subtidal fringe just be- Table 4). who provide floristic, geographic, low the lowest tide level and extend The warm-temperate flora of and ecological analyses, including a downward, but the majority are re- southern California does not offer detailed account of community stricted to depths of at least ten a mirror image of the pattern shown structure and productivity. Nearly meters. A few species occur as deep by the cold-temperate flora in which 60 species of California seaweeds, as 100 meters. Until the advent of species drop out progressively as the water temperature increases. Rather, it is largely confined to wa- Table 2. Conspicuous cold-temperate ters south of Point Conception, with seaweeds with mainland end-points between only a few species venturing farther the Monterey Peninsula and Goleta, Santa north. Those that do venture north- Barbara County, but also found on the ward are restricted to pockets of Channel Islands. relatively warm water, such as Stillwater Cove in Carmel and at Santa Cruz (see Table 5). At La Antithamnionella spirographidis (Schiffner) Wollaston Jolla and in the southern Channel Halymenia schizymenioides Hollenberg & Abbott Islands (San Clemente and Santa Mazzaella flaccida (Setchell & Gardner) Fredericq Catalina islands) there are several species that may be found in nearly Mazzaella splendens (Setchell & Gardner) Fredericq all subtropical and tropical waters. Neoptilota hypnoides (Harvey) Kylin As pointed out by Abbott and Neorhodomela oregona (Doty) Masuda Hollenberg (1976), the marine flora Porphyra variegata Setchell & Hus of southern California comprises Prasiola meridionalis Setchell & Gardner more species but less biomass than Urospora penicilliformis (Roth) Areschoug that of northern California, as is usually the case when a cold-water

24 FREMONTIA VOLUME 32:1, JANUARY 2004 scuba diving, several species were known only from plants cast ashore Table 3. Conspicuous cold-temperate after storms or pulled up in dredge seaweeds with mainland end-points in hauls. A few species have been ob- California between the Monterey served by divers, but are rarely if Peninsula and Goleta, Santa Barbara ever cast ashore, presumably disin- County, but also found on the Channel tegrating in situ. The geographic Islands and in northern Baja California. ranges of these subtidal species are erratic, in part depending on the availability of certain areas for sam- Bossiella californica (Decaisne) Silva pling. Many ranges encompass one Calliarthron tuberculosum (Postels & Ruprecht) Dawson or more of several areas favorable to scuba diving and dredging. These Callocolax fungiformis Kylin areas include Bowie Seamount Callophyllis obtusifolia J. Agardh (about 75 miles west of the Queen Chondracanthus corymbiferus (Kützing) Guiry Charlotte Islands, British Colum- Desmarestia latifrons K¸tzing bia), the Puget Sound (particularly Farlowia compressa J. Agardh Hein Bank in the San Juan Islands), Gloiopeltis furcata (Postels & Ruprecht) J. Agardh Cordell Bank (Marin County), the Farallon Islands (San Francisco Haplogloia andersonii (Farlow) Levring County), Monterey and Carmel Laminaria setchellii Silva submarine canyons and Schmieder Mastocarpus papillatus (C. Agardh) J. West Bank (Monterey County), Diablo Neorhodomela larix (Turner) Masuda Submarine Canyon (San Luis Peyssonneliopsis epiphytica Setchell & Lawson Obispo County), Scripps and La Schimmelmannia plumosa (Setchell) Abbott Jolla submarine canyons (San Di- ego County), and various diving and Scytosiphon dotyi Wynne dredging sites in the Channel Is- Soranthera ulvoidea Postels & Ruprecht lands and northern Baja California. including Tanner and Cortes banks. Kelp canopies, particularly those of Macrocystis pyrifera, provide a habi- explain. Part of the explanation may tat for a specialized algal flora to- be the deterioration of mainland gether with associated fishes and habitats. These endemics may have invertebrates. at one time been found on the main- A very few species of the Cali- land. The occurrence of several spe- fornia flora are insular endemics, cies of seaweeds that were found at occurring in the southern Channel San Pedro a hundred years ago, but Islands as well as on various Mexi- not in recent times, is documented can islands, especially Isla by specimens in the herbarium at Guadalupe and Rocas Alijos. The the University of California, Ber- high degree of endemism of the keley. Alternatively, these endemics marine biota of Isla Guadalupe is may be represented by small subtidal easily explained by the ecological coastal populations waiting to be isolation of this island resulting from discovered. Three conspicuous spe- its position to the west of the cold cies are currently considered strictly upwelled waters at the same lati- insular: Cystoseira neglecta Setchell tude on the Baja California penin- & Gardner and Sargassum palmeri sula. The contrast between the ma- Grunow (both brown algae) and an rine biota of Isla Guadalupe and undescribed species of Codium (a the mainland, which is only about green alga). 160 nautical miles to the east, is The Monterey Peninsula is re- Hesperophycus californicus, a conspicuous markably rich in seaweeds, both in brown warm-water rockweed, is only striking. The absence of these insu- rarely found north of Santa Cruz. Line lar endemics on the southern Cali- number of species and in luxuri- drawing from DeCew Guide, used with fornia mainland is more difficult to ance of growth. Its richness in part permission from P. Silva.

VOLUME 32:1, JANUARY 2004 FREMONTIA 25 is due to the equally remarkable ized factors that affect the distri- port of sand, and diversity of sub- variety of habitats in the area, so bution of seaweeds are always strate. An exemplary account of the that certain species find the lack of modified, to a significant degree in habitats and distribution of the a suitable habitat rather than higher certain places and at certain times, seaweeds and seagrasses of a par- temperatures a barrier to extend- by local conditions such as inshore ticular region is that by Stewart ing their range southward (see currents and countercurrents, lo- (1991), who shares in satisfying Table 1). calized upwelling, seasonal river detail her extensive long-term ob- In any given area, the general- and creek discharge, seasonal trans- servations of the marine flora of San Diego County.

Table 4. Conspicuous cold-temperate REFERENCES seaweeds with mainland end-points in California between the Monterey Peninsula Abbott, I.A. and G.J. Hollenberg. and Gaviota, Santa Barbara County, and 1976. Marine Algae of California. also found in northern Baja California Stanford University Press. but not in the Channel Islands. Stanford, CA. 827 pp. Dall, W.H. 1916. Checklist of the Re- cent Bivalve Mollusks (Pelecypoda) of Ahnfeltiopsis pacifica Silva & DeCew, ined. the Northwest Coast of America From the Polar Sea to San Diego, Califor- Blidingia minima (Naegeli) Kylin nia. Southwest Museum, Los An- Delesseria decipiens J. Agardh geles, CA. 44 pp. Desmarestia viridis (O.F. Mueller) Lamouroux Hickman, J.C. (ed.) 1993. The Jepson Porphyra gardneri (Smith & Hollenberg) Hawkes Manual: Higher Plants of California. Prionitis filiformis Kylin University of California Press. Ber- keley, CA. 1400 pp. Ptilota filicina J. Agardh 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 Salvador Table 5. Northern end-points of Island, Bahamas: Distribution, conspicuous warm temperate seaweeds. abundance, and primary productiv- - ity. Deep-Sea Research 33:881-892. Murray, S.N. and R.N. Bray. 1993. Santa Cruz Benthic macrophytes. In Ecology of Chondria decipiens Kylin the Southern California Bight, ed. M.D. Dailey, D.J. Reish, and J.W. Chondria nidifica Harvey Anderson, pp. 304-368. University Hesperophycus californicus Silva of California Press. Berkeley, CA. Silva, P.C. 1979. The benthic algal Monterey Peninsula flora of central San Francisco Bay. Acrosorium venulosum (Zanardini) Kylin In San Francisco Bay: The Urbanized Helminthora stricta Gardner Estuary, ed. T.J. Conomos, pp. 287- Laurencia pacifica Kylin 345. Pacific Division, American As- Laurencia subopposita (J. Agardh) Setchell sociation for the Advancement of Science, San Francisco, CA. San Luis Obispo County Stewart, J.G. 1991. Marine algae and Cambria seagrasses of San Diego County. Cali- Leptocladia binghamiae J. Agardh fornia Sea Grant College Report Shell Beach No. T-CSGCP-020. 197 pp. Cystoseira setchellii Gardner

Los Angeles County Paul C. Silva, University Herbarium, Redondo Beach UC, 1001 Valley Life Sciences Bldg. No. Laurencia snyderae Dawson 2465, Berkeley, CA 94720-2465. [email protected]

26 FREMONTIA VOLUME 32:1, JANUARY 2004 INTERTIDAL MONITORING OF CALIFORNIA SEAWEEDS by Kathleen Dickey

embers of the California needed to correlate changes in the and describing patterns over time. MNative Plant Society have seaweed flora and vegetation struc- These studies have been punctu- long been passionate ture with external forces, both ated and inspired by external events pioneers in documenting the iden- physical and anthropogenic. One such as oil spills, nuclear power plant tity, distribution, and abundance of prime lesson gleaned from over half development, sewage outfalls, and terrestrial plants. Botanists world- a century of intertidal monitoring other impacts of human population wide estimate that more than half is that the inherent patchiness in growth. of all the plants in the world have intertidal community organization It is challenging to census the now been identified. Marine plants makes extrapolation from observed marine environment consistently and systems have also been well changes to supposed causes chal- and over time, chiefly due to a lack studied, and more than 8,000 spe- lenging, and thus there is a great of funding and personnel for such a cies of marine macroalgae have been need for careful documentation. long-term commitment. It is also documented worldwide. In Califor- Intertidal monitoring studies fall often a physically demanding, or nia alone, more than 650 species into three general categories: 1) the even hazardous job—there is occa- of marine algae are known, but descriptive or “snap-shot” studies; sionally the possibility of being bit- there remains much to decipher 2) longer term studies, including ten by an angry elephant seal in the concerning patterns and changes in experiments testing questions that course of a count! A benefit of the community structure and species arose from the initial studies; and 3) focus on documention of intertidal richness. long-term, collaborative monitor- habitats and species has been the More research is especially ing projects dedicated to discerning renewed appreciation of and need

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

VOLUME 32:1, JANUARY 2004 FREMONTIA 27 Recognizing the differences in branching patterns between two species of Microcladia as well as the differences in habit is critical to an accurate census. Microcladia borealis (left, x2) is saxicolous (grows on rocks) whereas M. coulteri (right, x 1/2) is epiphytic (grows on other plants). Line drawings from the DeCew Guide, used with permission from P. Silva. for well-trained taxonomists, natu- intensively documented the rocky WAKE–UP CALLS ralists, and organismal biologists. intertidal inhabitants and their re- AND THEIR After all, it’s no use counting some- spective zones on the Monterey CONSEQUENCES thing that you can’t identify. Peninsula. One of the earliest for- mal intertidal censuses was con- In 1969 an oil platform blew ducted in a benchmark study in the out in the Santa Barbara Channel. A BRIEF HISTORY same area by W.G. Hewatt when The ensuing disaster prompted a he set up a permanent transect to series of studies, sponsored by the The first surveys done in the survey intertidal invertebrates at Minerals Management Services, in intertidal zone were primarily de- Hopkins Marine Station in 1931- order to obtain baseline data on the scriptive. Beginning in the 1930s 33. Unfortunately for the seaweed southern California coast. Suddenly the early studies (notably those done community, he didn’t include the surveys that had occurred in the by W.G. Hewatt, E. Yale Dawson, plants. Additional survey work was previous four decades had greatly P.K. Dayton, M.S. Foster, M.M. done by E. Yale Dawson, who moni- increased value. Littler, and S. Murray—for a re- tored 42 transects in southern Cali- In contrast to the Santa Barbara view of early work see Murray 2001) oil spill, which stirred both local fornia in 1963. were carefully executed, using clear and statewide interest, the 1989 In 1973, John Pearse and his and readily repeated methodology. Exxon Valdez disaster grabbed at- students, funded by Sea Grant and The locations of many of the early tention around the world, when local government agencies, con- studies were determined by access some eleven million gallons of oil to the tidepools and proximity to ducted a survey of a mudstone reef spilled into Prince William Sound, research stations. using meter-square plots along per- Alaska, fouling 1,300 miles of a here- E.F. Ricketts, author of the manent vertical transects (going tofore pristine intertidal region. seminal Between Pacific Tides and from the high tide to low tide zone). When it came time for Exxon to proprietor of a biological supply They established one transect fulfill their legal obligation to house (not to mention the inspira- beneath a sewage outfall that was remediate these Alaskan habitats tion for the main character in John closed down in 1976. They have after the disaster, quantitative data Steinbeck’s novel Cannery Row), continued to monitor these sites. were insufficient to establish what

28 FREMONTIA VOLUME 32:1, JANUARY 2004 had been lost. Unlike California, species. On the other hand, two available on the Internet. where there had been previous ef- northern snails increased in abun- We are now in the era of synthe- forts along the southern and cen- dance, but seemingly by migrating sis and long-term, carefully tral coast to collect intertidal data, to a lower spot in the intertidal. designed experiments to try to tease there simply wasn’t enough known Mean sea surface temperatures in- out the mechanism of change in about the species and communities creased about 3/4oC, during these the intertidal zone. The need for an in areas affected by the Alaskan spill. 60 years, with an increase of ap- accurate inventory of California’s in- As a consequence, Exxon is now proximately 3o C during July through tertidal flora is clear. With efforts of appealing the settlement for puni- August, the warmest months of the volunteers and professionals alike we tive damages. This case provided year. will hopefully be able to have a more an urgent wake-up call to those who In Santa Cruz, John Pearse and complete appreciation for the diver- monitor the marine environment. his students have continued to con- sity of this state’s seaweed flora, and If the acquisition of basic scientific duct yearly censuses, and after 30 for the beauty of the balance within knowledge was not a strong enough years have only recently noticed a and between intertidal ecosystems. incentive for establishing baseline recovery in diversity at the site of the data, then surely economics might sewage outfall. Initially, at the outfall provide a convincing argument. site, coralline algae were the over- REFERENCES whelmingly dominant species. In the late 1990s, signs of recovery at this Barry,J.P, C.H. Baxter, R.D. Sagarin, WHAT HAVE WE site, as measured by increased diver- and S.E. Gilman. 1995. Climate- LEARNED? sity, included the return of surfgrass, related, long-term faunal changes in soft-bodied invertebrates such as a California rocky intertidal com- munity. Science: 267:672-675. In some instances, scientists bryozoans, tunicates, and sponges, Dawson, E.Y. and M. Foster. 1982 and some of the delicate foliose red have documented considerable (2nd Edition). Seashore Plants of Cali- change. In other cases the results algae. This is an excellent example of fornia. UC Press. have been ambiguous. In follow-up a long-term study that has captured Murray, S.N., J. Goodson, A. Gerrard, surveys in the rocky intertidal of important changes resulting from al- and T. Luas. 2001. Long-term southern California localities, Steve leviating pollution stress. changes in rocky intertidal seaweed Murray and his students have ob- One important consequence of populations in urban southern Cali- served that the relative abundance the Exxon Valdez spill was new fund- fornia. J. Phycol. 37(3):37-38. of highly productive foliose algae ing of large-scale, publicly and pri- Pearse, J. Personal communication. has declined, to be replaced by turf- vately funded censuses. These include Raimondi, P.T., R.F. Ambrose, J.M. forming seaweeds, corallines, and efforts by the National Marine Sanc- Engle, and M. Wilson. 1999. Moni- toring of rocky intertidal resources crusts. The reasons for this may be tuaries, the Partnership for Interdis- along the central and southern Cali- ciplinary Studies of Coastal Oceans anthropogenic, as there are simply fornia mainland. Three-year report many more people coming to the (PISCO), the National Park Services for San Luis Obispo, Santa Barbara, southern shores than before. (including an All-Taxa Biological and Orange Counties (Fall 1995, In fact, it is estimated that over the Inventory at Tomales Bay in north- Spring 1998). Technical Report. US past three decades the population of ern California), and the Multi Agency Department of Interior, Minerals coastal counties in southern Rocky Intertidal Network (MA- Management Service, Pacific OCS California has increased by more RINE), a consortium of agencies and Region (MMS Cooperative Agree- than 50%. institutions in southern California. ment No. 14-35-0001-30761 with In 1995, 60 years after the In some cases, innovative inter- Southern California Educational Ini- Hewatt census of invertebrates at agency alliances have been forged tiative, Marine Science Institute, Uni- Hopkins Marine Station, Barry et between disparate groups, unified by versity of California, Santa Barbara). Ricketts, E.F. 1992. Between Pacific al. repeated the same work, using the desire to document and preserve Tides. Stanford University Press. the same transect and plots. They our marine heritage. Fishermen, stu- Stanford, CA. were surprised by what they found. dents, volunteers, and local residents Thom, R. M. and Widdowson, T. B. First, in 32 of the 45 invertebrate have been recruited to help univer- 1978. A resurvey of E. Yale Dawson’s species analyzed in detail, popula- sity and government scientists with 42 intertidal transects on the south- tion abundances changed signifi- their census work. Traditional tools ern California mainland after 15 years. cantly. Second, animals that were such as the plant press, transect, and Bull. S. Calif. Acad. Sci. 77:1-13. considered “southern” species were quadrat are now augmented with now more common, with a con- GPS and GIS technology, and data- Kathleen Dickey, PO Box 314, La Honda, comitant diminution in “northern” bases and images are often broadly CA 94020. [email protected]

VOLUME 32:1, JANUARY 2004 FREMONTIA 29 SEAWEED RESOURCES Compiled by Kathleen Dickey

If you are just beginning to learn represents a consortium of institu- about the wonders of seaweeds, the tions, state and federal agencies, and following list of reference books and private endeavors in southern Califor- websites will help you along your road nia conducting long-term intertidal of discovery. monitoring projects. Agencies include California State University, Fullerton, Minerals Management Service, Uni- BOOKS versity of Southern California, Uni- versity of California (Los Angeles, San Diego, Santa Barbara, and Santa Cruz Abbott, I.A. and G.J. Hollenberg. campuses), Regional Water Quality 1976. Marine Algae of California. Control Boards, PISCO (see website Stanford University Press. Stanford, below), California Department of Fish California, USA. and Game, the United States Navy, Dawson, E.Y. and M.S. Foster. the National Marine Sanctuary, and 1966. Seashore Plants of California. the National Park Service. University of California Press. Berke- www.biology.ucsd.edu/labs/roy/ ley, California. Revised 1986. CBRISC/summary.html. Conservation Druehl, L.D. 2000. Pacific Seaweeds: A Guide to the Common Seaweeds of the and Biodiversity of the Rocky Inter- West Coast. Harbour Publishing. Ma- tidal of Southern California (with the acronym CBRISC, pronounced deira Park, British Columbia, Canada. The red alga, Phycodrys setchellii. Line “SeaBrisk”). This site is aimed at assess- Lee, R.E. 1980. Phycology. Cam- drawing, to be posted soon to the DeCew bridge University Press. Cambridge, Website (see page 12), used with ing the health of Southern California’s England. permission from P. Silva. coastal marine environment. Mondragon J. and J. Mondragon. www.piscoweb.org. PISCO (Partner- 2003. Seaweeds of the Pacific Coast: Com- ship for Interdisciplinary Studies of www.coml.org/coml.htm. Census of mon Marine Algae from Alaska to Baja Coastal Oceans). PISCO is a large- Marine Life. This is a global network California. Sea Challengers. Monterey, scale marine research program that of researchers from more than 50 CA. examines near-shore marine systems. countries engaged in a ten-year ini- Ricketts, E. F. et al. 1985. Between The program is based at four univer- tiative (begun in 2000), with the goal Pacific Tides. Stanford University sities along the Pacific coast, and is of assessing and explaining the diver- Press. Stanford, CA, USA. funded by the Packard Foundation. sity, distribution, and abundance of www.farallones.org/education_page/ marine life in the oceans—past, studentprojects. html#rocky. Farallon WEBSITES present, and future. This endeavor is Marine Sanctuary Association funded by the Arthur P. Sloan Foun- (FMSA). Gulf of the Farallones dation. Two contributors to this site Seaweed information and National Marine Sanctuary, San are the Oceanic Biogeographic Infor- Francisco, California Educators work identification mation System(OBIS) and the Natu- with FMSA on two intertidal www.seaweed.ie. The National Uni- ral Geography in Shore Areas monitoring programs for students. versity of Ireland, Galway. This com- (NaGISA). limpets.noaa.gov/. Long term moni- prehensive website includes database toring program and experiential train- and images of seaweed from around Seaweed as food ing for students (LIMPETS). Olym- the world. www.seaweed.net. Mendocino Sea pic Coast, Cordell Bank, Gulf of the www.mbari.org/~conn/botany/. Vegetable Company, Mendocino Farallones, Monterey Bay, and Chan- Monterey Bay Aquarium Research County, California. nel Islands. Institute, Monterey, California. This www.loveseaweed.com. Rising Tide www.mbnms-simon.org/.The site includes student work from the Sea Vegetable Company, Mendocino Monterey Bay National Marine Sanc- Hopkins Marine Laboratory Marine County, California. tuary (MBNMS). One of 13 National Botany class. marine sanctuaries around the country, ucjeps.berkeley.edu/guide/. DeCew’s MBNMS has just posted an interactive Intertidal monitoring efforts Guide to the Seaweeds of British Co- database at their website, called the lumbia, Washington, Oregon, and www.marine.gov. Multi Agency Sanctuary Integrated Monitoring Northern California. See page 12. Rocky Intertidal Network. This site Network (SiMoN).

30 FREMONTIA VOLUME 32:1, JANUARY 2004 HARVESTING AND COLLECTING SEAWEEDS

California state Fish and Game 2) Some seaweeds are quite rare, regulations allows individuals to col- so if you find an individual plant of lect up to ten pounds of seaweed for any species, leave it alone. noncommercial uses. The best time to 3) Learn about the reproductive collect is just after a storm or when cycle of the seaweeds you are collect- the tide is going out, in order to en- ing, and what a fertile plant looks sure freshness. The best places to col- like. By not collecting fertile plants, lect drift seaweed are open ocean the population will be able to replen- beaches and rocky shores, away from ish itself. creeks or rivers. Seaweeds from bays 4) Do not assume that because a and inlets in California are best left plant is growing in the ocean it will be alone. There are no poisonous toxin-free. Some seaweeds selectively seaweeds, although one genus, uptake and concentrate minerals. Do Desmarestia, contains hydrosulfuric not collect seaweed near a city or near acid and is quite acrid. Others, such a stream or river that might carry ag- as sea lettuce (Ulva), are just too tough ricultural runoff into the ocean. As a to eat. general rule, bay waters (especially San Francisco Bay) should be off-limits.

GENERAL RULES REFERENCES 1) Before you pick any seaweed, make sure that there are many more Dawson, E.Y. and M. Foster. 1982. Sea- plants of the same species nearby. For shore Plants of California. University example, in the case of the sea palm of California Press. Berkeley, CA. The red alga, Ceramium californicum. (Postelsia), one study shows that there Druehl, L. 2001. Pacific Seaweeds. Line drawing, to be posted soon to the need to be at least 120 plants in a stand Harbour Publishing. Madeira Park, DeCew Website (see page 12), used with for it to reproduce. BC Canada. permission from P. Silva.

NOTES AND COMMENTS

CNPS FELLOWS NOT Beatrice Howitt, 1973 native to California, and is commonly UNDER–APPRECIATED Jim Griffin, 1993 found in a wide range of habitats. It Jake Sigg, 1997 is popular in the horticultural indus- Joan Stewart, 1999 try and is readily available from na- The January 2003 issue of Fremontia Randall Morgan, 2000 tive plant nurseries in California, Or- (Vol. 31(1):27) listed California Native In addition, there may be others egon, Washington, and British Co- Plant Society Fellows, but some of who should be included on this list. If lumbia. those honored with the title were in- you know if this is so and the date they As a host plant for the Sudden Oak advertently omitted from this list. Death pathogen, wood rose falls un- Apologies are given to these Fellows, became Fellows, please contact the Editor. der Phytophthora ramorum regulations, their friends, and families. Although a requiring inspection before being cer- Fellow’s name might have been over- tified for shipment from California looked, certainly their participation in SUDDEN OAK DEATH nurseries. For those planting natives the society is or has been greatly ap- in your garden, be careful to obtain preciated. UPDATES stock that is not infected with this Fellows names are listed below with pathogen. the date the title was received. In April of 2004, wood rose (Rosa For more information about Sud- Mary Ann Matthews, 1993 gymnocarpa) was added to the list of den Oak Death, go to the California Mary Rhyne, 1991 nearly 30 confirmed Phytophthora Oak Mortality Task Force (COMTF) Florence Marsh, 1991 ramorum host plants. Wood rose is website at: www.suddenoakdeath.org.

VOLUME 32:1, JANUARY 2004 FREMONTIA 31 LETTERS TO THE EDITOR

Weeds and Wildflowers considered the effect of plaintain and Aleins” should be ascribed to Doris (Volume 31, No. 2) weeds like it before. Ann Hoover, who sadly passed away Kathy Kramer recently. Doris was one of the origi- I wanted to say how much I enjoyed East Bay Chapter nal weed warriors in the LA Chapter, Jake Sigg’s article on weeds, and also and these comments appeared first, if the article by Marjorie Schmidt! It Weed Reference Correction I remember correctly, in the local would be great if Fremontia ran an ar- (Volume 31, No. 4:34) Toyon. ticle of historical quotes in every is- Michael O’Brien sue. Jake’s article was so well-written In the “Selected Articles on Weeds,” Los Angeles/Santa Monica and thought-provoking—I had not the reference to “Control of the Chapter

BOOK REVIEW

Pacific Seaweeds: A Guide to the corporates taxonomy based on exter- the fascinating stories and asides Common Seaweeds of the West nal morphology (for example, crusts, that range from the intriguing to Coast, by Louis Druehl. 2000. globular, small flat blades, cylinders) the preposterous. For example, he de- Harbour Press, Madeira Park, British with a small “thumbnail” dichoto- scribes the sea palm, Postelsia Columbia, Canada. Softcover, $17.95. mous key. The photographs and line palmataeformia, as being “severely Louis Druehl has been studying and drawings are very helpful in the field. attached to a rock.” Additionally, he teaching about seaweeds for more than As one must in any small book, includes many simple recipes, some 30 years, and recently started farming Druehl focuses on some 100 of the complete with verse. Altogether, this kelps in British Columbia. In this book more common seaweeds, perhaps is a satisfying introductory book to he presents information and fascinat- with a disproportionate emphasis on accompany you in forays to the ing lore about seaweeds aimed at a his specialty, the “browns.” tidepools. It includes a helpful glos- general, interested audience. He in- My favorite parts of this book are sary and list of references.

RECIPE FOR MARINATED ALARIA The kelp, Alaria marginata. Line drawing from DeCew Guide, used Thanks to Louis Druehl and Rae Hopkins, with permission from P. Silva. Canadian Kelp Resources, Ltd. Ingredients 1 cup dried Alaria, cut in strips (use scissors) 1/2 cup rice vinegar 3 Tbsp soy sauce 1 Tbsp sugar 1 tsp salt Optional and delicious additions Thinly sliced cucumber Finely grated carrots Sliced green onions

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

32 FREMONTIA VOLUME 32:1, JANUARY 2004 Please Join Today! CNPS member gifts allows us to promote and protect California’s native plants and their habitats. Gifts are tax-deductible minus the $12 of the total gift which goes toward publication of Fremontia and the CNPS Bulletin.

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MATERIALS FOR PUBLICATION CLASSIFIED ADS Members and others are in- vited to submit material for pub- Classified ad rate: $1.00 per word, notecards, books, fruit crate labels, hand lication in Fremontia. Instruc- minimum $15; payment in advance. towels, and T-shirts focusing on tions for contributors can be Address advertising inquiries and copy to: California native plants from Sierra found on the CNPS website, CNPS, 2707 K Street, Suite 1, Sacra- Nature Prints. Animal puppets, too. Visit mento, CA 95816-5113. (916) 447-2677 www.sierranatureprints.com www.cnps.org, or can be re- or fax (916) 447-2727. quested from Fremontia Editor, NURSERIES AND SEEDS Linda Ann Vorobik, vorobik@rock PUBLICATIONS Telos Rare Bulbs. Bulbs for your gar- island.com, or c/o University and Flora & Fauna Books, 121 First Avenue den, restoration projects, landscaping. Jepson Herbaria, 1001 Valley South, Seattle WA 98104, Tel. (206) Many Calif. native species, including Life Sciences Bldg. #2465, Uni- 623-4727, Fax (206) 623-2001, ffbooks Calochortus, Fritillaria, Brodiaea rela- @blarg.net, Specializing in Botany, tives, Erythronium. Catalog $3.00. Free versity of California, Berkeley, shipping in USA. P.O. Box 4978, Arcata, Gardening, Birding, and Ecology, both CA 94720-2465. CA 95518. www.telosrarebulbs.com. new and out-of-print. We carry a large inventory of floras, keys, and field guides ECOLOGICAL RESTORATION for the west coast and worldwide. A FREMONTIA EDITORIAL large selection of our inventory is now Native California grassland and desert ADVISORY BOARD available on the web: www.abebooks.com/ ecological restoration standards and home/FFBOOK/. costs, plus pictures showing results. Ann Bradley, Susan D’Alcamo- www.ecoseeds. com/standards.html. ART Potter, Ellen Dean, Kathleen SERVICES Dickey, Phyllis M. Faber, Bart Notecards, Prints, and Originals. Visit O’Brien, John Sawyer, Jim www.VorobikBotanicArt.com. PO Box Nature landscape design. Landscape 866, Lopez Island, WA 98261 Design that celebrates the rich heritage Shevock, Teresa Sholars, Nevin of California’s native flora. Duber Land- Smith, Dieter Wilken, John Botanical prints, note cards, postcards, scape Design, CA license #4316. (510) Willoughby, Darrell Wright Plants of the Lewis and Clark Expedition 524-8665.

VOLUME 32:2, APRIL 2004 FROM THE EDITOR

he seashore provides a con- Paul C. Silva and Kathy Ann Miller, Dickey has provided, or try the recipe T trasting experience to that have through their combined efforts from the book review on page 32. which we find with walks to offered several perspectives from This issue begins my fourth year as meadow or mountain. The smell of which to consider algae. Silva relates Editor of Fremontia, and during that salt, view of ocean, and sound of wind stories of their discovery and how they time my goal has been to provide a and wave all combine to make a visit are harvested for useful products. He variety of articles that are backed by to the intertidal unique. This special also carefully describes how currents academic research and yet are inter- issue of Fremontia leads us into the along our coast contribute to the great esting and available to a broad audi- world of seaweeds, and provides pre- diversity of California seaweeds. ence—with articles ranging from wild- liminary information along with a few Miller informs us of invasive seaweeds flower appreciation and horticulture, tools of understanding that may make that we, as stewards to natural envi- through ecology and taxonomy, to your next visit there as joyful as any ronments, should be on the lookout advocacy. I wish to make these issues meadow stroll. And if you glory in di- for. Dickey provides background in- the best they can be for our society, versity and detail, you will be delighted formation on seaweed research as well and welcome all letters that respond by this group of plants that differ so as how seaweeds are monitored, and to individual articles or to the general from our more familiar flowers of why it is important to continue study- scope of our California Native Plant forest and field. ing them. And if you are inspired to Society journal, Fremontia. The Convening Editor, Kathleen learn more, look to the back pages for Linda Ann Vorobik, Dickey, along with the contributors a list of Web and other resources that Editor CONTRIBUTORS

Tom DeCew received his MS at Humboldt State University and his PhD at UC Berkeley. During his studies he canvassed West Coast herbaria to compile phenology and distribution

data and searched the macroalgal literature for studies pertain-

Address Service Requested Service Address

Sacramento, CA 95816-5113 CA Sacramento, 2707 K Street, Suite 1 Suite Street, K 2707 ing to West Coast seaweeds. The fruits of his labor resulted in Society Plant Native California “The Book,” a bibliography to the biology of common north- east Pacific seaweeds with line drawings and tabular displays of latitudinal distribution and reproductive phenology. It was in- tended to complement Marine Algae of California by I.A. Abbott and 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, UC Berkeley. She received her MS at UCSC. She has worked as a consultant 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 of Southern California’s Wrigley Marine Science Center on Santa Catalina Island. She earned her BA and PhD degrees in Botany at the University of California at Berkeley. She specializes in seaweeds 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-

U.S. Postage U.S.

Nonprofit Org. Nonprofit Permit # 3729 # Permit

nia Channel Islands. CA Oakland, PAID Paul C. Silva is a native son of California. He received his BA at the University of Southern California, his MA at Stanford University, and his PhD at the University of California, Berke- ley. He has been on the staff of the University Herbarium since 1960 and is considered a world authority on seaweeds.

VOLUME 32:1, JANUARY 2004