An Introduction to the Seaweeds of British Columbia by Colin Bates, Dept of Botany, University of British Columbia. Adapted from an account originally published on E-Flora BC website. All photos by Colin Bates.

any people are unaware can be large or small, helpful or endosymbiotic events gave rise to that British Columbia is a harmful, and believe it or not, red and green algae (e.g., Moreira et Mveritable hot spot of sea- stunningly beautiful and head- al. 2000). It is generally accepted weed biodiversity. At last count, scratchingly interesting. that there were multiple secondary there were approximately 650 endosymbiotic events. With the Unlike plants and animals, the algae macroalgal species found in waters advent of molecular tools, algal do not arise from a single common of the Pacific Northwest (Scagel et systematics is in the midst of a ancestor (a condition referred to as al. 1989, Gabrielson et al. 2000). Of substantial reworking of these and monophyly). Instead, they are a these, 530 are known to occur in other questions. group composed of many lineages BC. (group composed of many lineages; Molecular tools have also provided Why are so many people unaware of i.e. polyphyletic (Graham & Wilcox the opportunity to examine relation- the diversity of seaweeds that in- 2000). Throughout the history of ships between algae and land plants habit British Columbia’s waters and algal , various characters (Bhattacharya & Medlin 1998). shores? Many initial encounters with have been invoked to describe this Mounting evidence suggests that seaweeds can be unpleasant, disparate group: simple bodied land plants are derived from an especially if you are wading into the organisms, reproductive propagules ancestor of the green algal order ocean for a swim, or trying to main- lacking well-developed structures as Charales (Mishler & Churchill 1985, tain your footing while scrambling in flowers or cones, mostly aquatic, Surek et al. 1994). Land plants are on the rocks. As well, detached and generally photosynthetic. Yet, excluded from algae because they do seaweeds end up washing ashore because the algae are polyphyletic, not conform to the character set and amassing into smelly decom- arising from various ancient described above; these plants have posing piles. Seaweeds generally get ancestors, there are myriad complex bodies, typically with a bad reputation in the press (if any exceptions to these characters as flowers or cones, and are normally at all). Books like “Killer Algae” well as many non-algal taxa that terrestrial. (Meinesz, 1999), which describes possess them. The study of algae is called phy- the invasive nature of a green The tie that binds all algae together cology, from the Greek root phykos seaweed species called Caulerpa is a concept known as the ‘serial meaning algae. Algae are a vast taxifolia, and talk of ‘Harmful Algal endosymbiosis hypothesis’ group including many classes of Blooms” (a microalgal phenom- (Bermudes & Margulis 1985, unicellular and multicellular enon) do not help. All of these Bhattacharya 2000), which suggests organisms. Below, I focus on the factors contribute to the idea that that the commonality among all multicellular macrophytes, and in seaweeds are a nuisance. Yet, it is algae is that, at one or more points in particular, the benthic marine macro- my experience that seaweeds are their lineage, they acquired photo- phytes that most refer to as the most certainly not a nuisance, and synthetic abilities through either the seaweeds. instead are beautiful organisms with uptake of a photoautotrophic bac- fascinating natural history, diverse Seaweeds are a macroscopic subset terium (known as a primary endo- economic uses, and essential of marine algae (as opposed to the symbiotic event), or through the ecological roles. microscopic subset known as phyto- uptake of a heterotroph that had plankton). The west coast of British Seaweeds are a subset of a larger previously taken up a photoauto- Columbia has a diverse flora and group known as the algae. When trophic bacterium (a secondary arguably is home to the widest most people think of algae, they endosymbiotic event). A current hot variety of kelp (Order Laminariales) picture a green scum on the side of a topic in algal phylogenetics is in the world. pond or fish tank, but in reality algae determining if one or two primary

4 • menziesia The seaweeds are split into three Cladophora is a genus represented major groups, originally delineated by up to eight species in the Pacific by their pigment complement (and Northwest. Cladophora is hence color): the green algae superficially similar to Acrosiphonia (Division Chlorophyta), the brown but is usually shorter and lacks algae (Class Phaeophyceae, hooked branches. Patches of Division Ochrophyta), and the red Cladophora look instead like tufts of algae (Division Rhodophyta). I will moss on exposed rocks or spreadout refer to them henceforth as the filaments in tidepools. greens, browns and reds. Codium is locally represented by Codium setchellii three species, but only two are Green seaweeds Ulva is represented by eleven species common: C. setchellii and C. fragile. The greens are often a brilliant in British Columbia. Traditionally, The cells of Codium are grassy green, although a few such Ulva was recognized as a thin ‘coenocytic,’ meaning that they lack as Codium setchellii are so dark that distromatic (two-cell layer) green cross-walls in their cells. As a result, they appear almost black. The blade, which is fast growing and a an individual of C. fragile meas- bright green stems from a lack of weedy. You may confuse Ulva with uring up to 25 cm can be composed accessory pigments to mask the numerous other less-common green of a single multi-nucleate cell! chlorophyll a pigments used in blades, such as Ulvaria, . There are many Economically, kelps and rockweeds Monostroma, or Kornmannia. These are harvested and used in their raw freshwater green algae, but some three genera are monostromatic (one- classes of greens are almost form for food products, cosmetic cell layer) blades. Recent molecular and luxury spa items, and fertilizers. exclusively marine (e.g. evidence (Hayden et al. 2003) Ulvophyceae, which contains the In addition, through processing, their demonstrated that Ulva also cell wall components (alginic acids genera Ulva and Acrosiphonia). encompasses the genus previously Green seaweeds come in a wide and other polysaccharides) can be called Enteromorpha. Traditionally, it removed and used as emulsifiers, variety of morphologies, ranging was separated from Ulva because of from unicells and filaments to anticoagulants, and in the production its tubular growth form. As a result, of textiles and rubbers. blades and fleshy forms. species previously known as In general, the greens are Enteromorpha are now known as Ecologically, kelps and rockweeds encountered in the intertidal zone. Ulva, (e.g. Enteromorpha intestinalis are major habitat providers and The diversity of greens in the = Ulva intestinalis). nursery environments for fish, Pacific Northwest includes 117 taxa invertebrates, and for other algae. Acrosiphonia is a branched Their extensive biomass provides a in 51 genera (Gabrielson et al. filamentous green. The locally 2000). Common genera of green large amount of primary produc- abundant species Acrosiphonia tivity, carbon and oxygen to near- algae in British Columbia include: coalita grows in clumps resembling Ulva (including Enteromorpha), shore food webs. Further, nearshore green dreadlocks due to the presence kelp beds protect shorelines from Acrosiphonia, Cladophora, of tiny hooked branches. Codium, and Prasiola. erosion by decreasing the impacts of Consequently, its common name is water motion. “Witches Hair.” Interestingly, for part of The fucoids are represented in its life history, British Columbia by four genera: Acrosiphonia is endo- Fucus, Sargassum, Cystoseira, and phytic inside certain red Pelvetiopsis. Fucus is likely the algal species (Sussmann most conspicuous seaweed in British & DeWreede 2001), Columbia, represented by two such as Mazzaella species: Fucus distichus subsp. splendens and the evanescens (also known as Fucus ‘Petrocelis’ phase of gardneri) and Fucus spiralis. (see the Sargassum is represented by one Ulva sp. section about red introduced species, S. muticum, seaweeds below). which is suspected of entering

autumn 2004 • 5 British Columbia prior to 1940 as a sulfuric acid (pH approximates 2; byproduct of oyster spat importation the same as your stomach!) when from Japan (Scagel 1956). To keep disturbed or exposed to air for too oyster spat moist and cool during long. You may see it digesting transport, S. muticum was used as itself after being swept ashore after wrap, and presumably was dumped a storm. If collecting, make sure into British Columbian waters. this one goes into its own bag with Cystoseira is a less common alga that plenty of seawater. resembles Sargassum, and are economically Pelvetiopsis limitata, which looks important. Directly or indirectly much like juvenile Fucus but lacks a (through extraction), red algae are midrib, is common in the upper part of daily life for many North intertidal zone of wave-exposed Americans as food, for compounds British Columbian shores. made with their cell wall compon- The diversity of kelp in British ents, and for their potential as Columbia is nearly unparalleled. Over sources of natural chemicals. 30 species of these beautiful brown Food: Many red algae are eaten. algae can be found in most outer coast The most familiar is ‘nori’ rocky habitats, ranging from the mid ( yezoensis), used to Postelsia palmaeformis intertidal to the subtidal zones. The make sushi rolls. Nori is farmed most conspicuous are the kelp forests Common reds along the BC coast extensively in Asia and was grown composed of the giant kelp, Macro- include: Mazzaella splendens, briefly in BC in the 1980s. Local cystis integrifolia, and bull kelp, Porphyra spp., Chondracanthus species are harvested here for Nereocystis luetkeana. Among the exasperatus, Microcladia coulteri, personal use by First Nations most unique, the sea palm, Postelsia Mastocarpus papillatus, Prionitis (Turner 2003) and others. Species palmaeformis, is found on the most lanceolata, and various coralline of Palmaria, related to the dulse of wave-battered shores. Its rarity and algae. the Maritimes, are also harvested palm tree-like appearance make it a for personal consumption. treat to view. Mazzaella splendens is a common red blade in British Columbia, Cell wall components: Red algal recognized by its iridescent sheen in cell wall components, in particular the lower intertidal zone and tide- agars and carrageenans, are used in pools. This sheen is caused by the cosmetics, food preparations, and same phenomenon as the iridescence biomedical and biotechnology caused by oil on water: multiple research. Ice cream contains layers (of cuticle in the case of carrageenan to emulsify milk and Mazzaella) cause light to be water. Toothpaste has red algal differentially refracted, giving the byproducts included to keep the multiple colors of reflected light. calcium carbonate component mixed with the aqueous com- Chondracanthus exasperatus is ponent. So, it seems red algal by- commonly known as the “Turkish Nereocystis sp. products play an integral role in Towel.” Its red blade is distin- causing and preventing tooth decay! Other interesting on BC guished by its very bumpy (papil- late) texture. It grows in the lower shores include the bizarrely Natural Chemicals: Red algal intertidal zone, though blades can be convoluted and brain-like Leathesia species are being intensively washed up on the beach after a difformis, ‘sausage-weed,’ Scytosipon surveyed for medicinal properties storm. As implied by name, this is lomentarius, and the bracket fungus- and antiviral agents. Evidence excellent for use in the shower as a like Ralfsia fungiformis. Also, a suggests that chemicals from red natural exfoliating cloth! particularly interesting genus is algae can help fight viral infections Desmarestia, the acid seaweeds. Some (Richards et al. 1978), including Microcladia coulteri is a finely species have an anti-herbivory herpes (Dieg et al. 1974) and HIV branched and beautiful species. It is adaptation that involves the release of (Luescher-Mattli 2003). generally found as an epiphyte (a

6 • menziesia plant growing on another plant) on herbivory. This group incorporates I hope this summary convinces you Mazzaella splendens and Chondra- calcium carbonate into its cell walls, that benthic marine algae (seaweeds) canthus exasperatus in southern BC. resulting in a very hard skeleton. are important, interesting, and Both crustose forms and upright integral components of the coastal Mastocarpus papillatus is a common geniculate forms can be found in the ecosystem. True, there are some alga with two very different growth Northeast Pacific and are nuisance algae, but the majority are forms. The upright form, known identifiable by their pink color. unimposing, simply ‘going with the commonly as the ‘Turkish flow’ of the waxing and waning Washcloth,’ is associated with the The reds are among the most beaut- tides. Next time you are on the rocky genus name Mastocarpus. It is a iful algae, but usually the most shore, take some time to discover small, typically bifurcating, dark underappreciated because of com- these natural wonders! reddish brown blade with papillae. plex life cycles and the difficulty of However, the alternate phase is a identification. However, I assure you Acknowledgements: This is dedicated to blackish crust that looks like tar on that if you spend some time getting the professors of phycology who taught the rocks. For a long time this phase to know the reds, they will become me what I know about seaweeds: Louis your favorite group of Druehl, Gary Saunders, Thierry Chopin, seaweeds! and Robert DeWreede. This article was greatly improved by comments from There are numerous Sandra Lindstrom. marine macrophytes References and other marine Bermudes D, and L Margulis (1985) organisms with photo- Symbiosis as a Mechanism of Evolution: synthetic endosym- Status of the Symbiosis Theory. Symbiosis bionts that are not 1: 101-124. considered seaweeds. Bhattacharya D (2000) A molecular phylogenetic perspective on the primary Seagrasses (Zostera and secondary endosymbiotic origins of spp.) and surfgrasses algal plastids. Algae 15:1-5. (Phyllospadix spp.). Bhattacharya D, and L Medlin (1998) These genera are Algal phylogeny and the origin of land angiosperms that have plants. Plant Physiol. 116: 9-15. moved into the marine Dieg et al.(1974) “Inhibition of herpes environment. Although virus replication by marine algae Microcladia coulteri they are photosynthetic extracts,” Anitimicrb. Ag. Chemother. marine macrophytes, 6:524-525. was thought to belong to a totally they reproduce using flowers and Gabrielson PW, Widdowson TB, separate species known as Petrocelis seeds. Both genera are important Lindstrom SC, Hawkes MW, and Scagel, franciscana. Culture experiments marine plants and hosts to various RF (2000) Keys to the Benthic Marine demonstrated that the two phases epiphytic marine algae, but are not Algae and Seagrasses of British Columbia, were different parts of the life cycle seaweeds. Southeast , and of one species. Hence, we refer to Oregon. Phycological Contribution #5, the crust as the ‘Petrocelis phase’ of Marine lichens (e.g. Verrucaria University of British Columbia, the Mastocarpus life history. spp.). Although lichens are a Department of Botany. 189 pp. symbiosis between cyanobacteria Graham L and Wilcox (2000) Algae. Prionitis lanceolata is known as (an alga) and a fungus, and marine Prentice Hall. ‘bleach weed.’ It usually grows in lichens do exist, but are not Hayden HS, Blomster J, Maggs C, Silva the lower intertidal zone, though it considered seaweeds. PC, Stanhope MJ, and Waaland JR can be found in tidepools higher on (2003) Linnaeus was right all along: Ulva shore. Its common name stems from Corals and Sea Anemones. Corals are actually colonial animals, though and Enteromorpha are not distinct genera. the scent of bromine the plants emit. Eur. J. Phycol. 38: 277-294. Presumably, the bromine acts as a deriving some energy from algal Luescher-Mattli M (2003) Algae, A chemical defense to herbivory. symbionts. Sea anemones also have endosymbiotic algae (which can Possible Source for New Drugs in the The coralline algae (Order Coral- affect the colour of the anemone), Treatment of HIV and Other Viral Diseases. Current Medicinal Chemistry - linales) have a unique adaptation to but sea anemones are also animals. Anti-Infective Agents 2: 219-225.

autumn 2004 • 7 Meinesz A (1999) Killer Algae. University Papers 1(4): 49-58. Sussmann AV and RE DeWreede (2001) of Chicago Press. 376 pp. Scagel RF, Gabrielson PW, Garbary DJ, Life history of Acrosiphonia (Codiolales, Mishler BD & Churchill SP (1985) Golden l, Hawkes MW, Lindstrom SC, Chlorophyta) in southwestern British Transition to a land flora: phylogenetic Oliveira JC, and Widdowson TB (1989) Columbia, Canada. American Journal of relationships of the green algae and A Synopsis of the Benthic Marine Algae Botany 88: 1535-1544. bryophytes. Cladistics 1: 305-328. of British Columbia, Southeast Alaska, Turner NJ (2003) The ethnobotany of Moreira H, Le Guyadev, and H Phillipe Washington and Oregon. University of (Porphyra abbottae and (2000) The origin of red algae and the British Columbia, Vancouver, related species; Rhodophyta: Bangiales) evolution of chloroplasts. Nature 405: Phycological Contribution No. 3, 532 and its use by First Nations on the Pacific 69-72. pp. Reprinted 1993 with minor revisions. coast of Canada. Can. J. Bot. 81: 283- 293. Richards et al. (1978) Antiviral activity of Surek B, U Beemelmenns, M Melkonian, extracts from marine algae. Antimicrob. and D Bhattacharya (1994). Ribosomal Agents Chemother. 14: 24-30. RNA sequence comparisons demonstrate an evolutionary relationship between Scagel RF (1956). Introduction of a Zygnematales and charophytes. Pl. Syst. Japanese alga, Sargassum muticum, into Evol. 191:171-181. the Northeast Pacific. Fisheries Research Indian Plum in the Salmon River Valley by Rose-Marie Silkens

emleria cerasiformis doesn’t I chose a spot in open woodland, last four years they have begun to occur in the wild on mixed trees and shrubs, quite grow more quickly and are now Onorthern Vancouver Island. sheltered from both winter south- about four feet high and as wide or When I lived in Victoria I enjoyed easters and cold, dry northwesterlies. wider. The exception is the plant in seeing its unbelievably-green early The soil is reasonably good as we the more open site, which has taken leaves along roadsides in Saanich, are on the old floodplain of the on a tree-like habit, and is well over but never noticed it north of that Salmon River, alluvial deposits six feet tall. It was outstripping its rarefied environment. It appears topped up with a few decades of neigh-bour, the ash, until two years sporadically as far north as Comox, humus-rich duff from alder, fir and ago. That tree took its time to but not in my current home in hemlock. One plant went into a more establish, but has come along very Sayward, 50 miles north of Camp- open site, at the edge of this well also. bell River. That city is at the end of woodland. I had purchased a native There is a note in Pojar and Georgia Strait, whose wide sweep Oregon ash, Fraxinus latifolia, on MacKinnon (1994) that native has a moderating effect on the east the same trip and planted it nearby. people reportedly ate Indian plum coast of Vancouver Island. Sayward I was surprised when every plant found near the head of Knight Inlet, is close to the narrow Johnstone survived that first season, though “although O. cerasiformis has not Strait, which sends bitingly cold growth was very modest. There were been recorded there.” It’s interesting winds well into spring. I’ve assumed no flowers for the first five or six to remember that Knight Inlet is that the very early blossoms on years. The first plant to bloom, in very near us, although the head is Oemleria would lose any viability to 2000, was in the open site, and by somewhat further north. February and March frosts. the next two years all were blooming My hope now is that fruit from my Nonetheless, when we saw a small by February. The only plant to set imported oemleria will produce patch of Oemleria in the Cowichan fruit is the more exposed one. Its some seedlings here and there. Valley, clearly destined for destruc- fruit set was reasonably generous Certainly, it wouldn’t be hard to spot tion, I decided to try taking some this year, despite a cold, damaging that spectacular spring green when back to Sayward. They were grow- early spring. Many normally-hardy the surrounding world is still dark. ing at the edge of a pushout in the cultivated plants in my garden suf- path of road construction, so that fered significant damage in March. Reference: Pojar, J. and A. MacKinnon weekend would have been their last. (Eds.) 1994. Plants of Coastal British I have never cultivated or interfered It was March 1996, and cold; I took Columbia including Wahington, Oregon with the transplanted Oemleria, a few cuttings before planting out & Alaska. Lone Pine Publishing, other than occasionally to remove my new treasures. Vancouver. 526 pp. encroaching thimbleberries. In the

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