Phaeophyta and Rhodophyta Lecture 9: Kelp Habitats

Fan-Shaped Phylogenetic Tree

Algal pigments

Algal pigments

Pigment Molecules Fucoxanthin Phycobilins

Accessory Pigments “Chromatic adaptation” Action spectra Chlorophylls 440, 670nm) Fucoxanthin (440-460nm) Phycobilins – phycoerythrin(540nm) & phycocyanin(620nm)

Div Phaeophyta Taxonomically maybe placed as a Class Phaeophyceae in Div Chromophyta/ Chrysophyta/ Heterokontophyta!!! Phaeophyta (brown) – 997 spp (almost exclusively marine. Chl a + c, B-carotene, fucoxanthin, neofucoxanthin, carotenoids Size: filamentous(mm) to kelps(m) 14 Orders: e.g. Ectocarpales, Sporochnales, Dicytotales, Fucales, Laminariales Temperate, few tropicals

Sargassum Padina Dictyota Laminaria

Div Rhodophyta Rhodophyta (red) – 4000-6000 spp 97% are marine No flagellae, pit connections (in Florideophyc) Chl a, a + B-carotene, xanthophylls, phycobiliproteins (red) Filamentous (mm) to corticated (dm), crustose (CaCO3) 2 Classes (Dixon 1973): Bangiophyceae (Porphyra) + Florideophyceae (Polysiphonia, Ceramium, Chondrus) Temperate and tropical.

Caloglossa Laurencia Polysiphonia Kallymenia Falkenbergia

Phaeophyte taxonomy Single class: Phaeophyceae. Almost exclusively marine, only 5-6 genera FW. Primarily temperate, some tropicals Lithophytes – requires hard substrate Sargassum are free-floating – Sargasso sea 14 Orders, e.g. Ectocarpales, Fucales, Laminariales

Thallus organization Filamentous – Ectocarpus Multiseriate filaments – Sphacelaria Crustose - Ralfsia Parenchymatous – Dictyota, Padina Morphologically differentiate – Laminaria, Fucus Taxonomy based on reproductive structures and life cycles

THALLUS COARSLEY - BRANCHED FILAMENT SHEET ENCRUSTING JOINTED - CALCAREOUS THICK - LEATHERY

Macroalgae - Phaeophyta

Class Phaeophyceae taxonomy Ectocarpus: Unilocular vs Pleurilocular sporangia Dawes, pg 135

Or. Ectocarpales Pilayella Ectocarpus

Or. Sphacelariales Sphacelaria

Or. Ralfsiales Ralfsia verrucosa

Or. Dictyotales – 16 genera Dicytopteris sp – mid rib Dictyota dichotoma

Or. Dictyotales Stypopodium – fish deterrant Padina - calcified

Or. Dictyotales – Lobophora Crust form Grazing pressure Decumbent form Ruffled form

Or. Chordariales Cladosiphon

Or. Sporochnales - GoM S. apodus S. moorei S. radiciformis

Or. Desmarestiales Sulfuric acid Desmarestia ligulata

Or. Scytosiphonales F. Chnoosporaceae F. Scytosiphonaceae Colpomenia sinuosa (sea corn-flakes) Hydroclathrus Scytosiphon lomentaria

Or. Laminariales “Kelps” Northern hemisphere 4 families: Chordaceae Laminariaceae (Laminaria, Agarum) Lessoniaceae (Macrocystis, Postelsia) Alariaceae (Alaria, Egragia) *Oceanographic conditions determine these 3 factors Next to coral reefs, considered largest biological structure in the oceans Useful species, common names, some of their uses? SpeciesCommon nameUses acanthopeltis agar Ahnfeltia plicata in USSR agar Anaebaena-Azolla symbiosis "green manure" Inoculation of 0.2 kg of Azolla per hectare is equivalent to 30 kg/hectare of commercial nitrogen fertilizer. Not as popular now because we have found that Azolla growth is limited by phosphate, which is non-renewable. Next break is low-phosphorus requiring Azolla Ascophyllum nodosum (bladder wrack) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Calcareous algae to reduce soil acidity. Caulerpa racemosa (Nama) in Fiji food Chlamydomonas and other soil algae soil amendment by making mucilage that loosen compacted soil. Chlorella only microscopic green used commercially Chondrus crispus (Irish Moss) agar, carrageenan Codium geppii (Sagati) in Fiji food Diatom fossils (diatomaceous earth, diatomite) flea killers, embalming compounds, enamel polishes, diatomaceous bricks, to filter sewage in UK from 1976, to filter sugar cane liquors from 1914; Nobel stabilized nitroglycerine. Durvillaea (Australia & Chile) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Ecklonia alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Eucheuma in Philippines carrageenan Furcillaria fastigiata in Denmark agar Gelidium amansii in Japan agar Gelidium arborescens in USA agar Gelidium cartillagineum in USA agar Gelidium latifolium in Ireland. agar Gelidium nudifrons in USA. agar Gelidium pulchellum Ireland agar Gigartina clavifera in New Zealand carrageenan Gigartina mamillosa (very similar and known as Irish Moss). carrageenan Gigartina undulata in New Zealand carrageenan Gracilaria (Lumicevata) in Fiji food Hypnea (Lumiwawa) in Fiji food Kelp and mixed seaweeds liquid fertilizers, soil amendment Wayne and Wanda! rich in potassium and nitrogen but low in phosphate. Free of terrestrial weeds and fungi. Kelps (Kombu) vegetables in Japan and China Laminaria (kelp - English; kombu - Japan and China) Food. Maybe 75% of their dry bulk is indigestible Laminaria digitata in UK and Norway alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Laminaria cloustoni (oar weed) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Macrocystis pyrifera (kelp - USA) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Mastocarpus stellata in British Isles. agar Phyllophora nervosa in USSR. agar Porphyra (laver - England and USA; luche - Chile; karengo - Maori; nori - Japan; slack - Scotland; sloke - Ireland) Food - Maybe 75% of their dry bulk is indigestible. Toasted and used to wrap sushi North Pacific. Cultivated for centuries in Japan, Korea and China Pterocladia agar Spirulina food protein - blue-green used for protein food in old (Lake Chad) and new world (Aztecs before conquistadores) Ulva food - eaten as a green vegetable Undaria pinnatifida (wakame) food - Maybe 75% of their dry bulk is indigestible http://seaweed.ucg.ie/Algae/laminaria.html Laminaria digitata

Or. Fucales Southern Hemisphere Wrack or Rockweed 4 Families Fucaceae (Fucus, Ascophyllum) Sargassaceae (Sargassum, Turbinaria) Cystoseiraceae Hormosiraceae (Hormosira) *Oceanographic conditions determine these 3 factors Next to coral reefs, considered largest biological structure in the oceans Useful species, common names, some of their uses? SpeciesCommon nameUses acanthopeltis agar Ahnfeltia plicata in USSR agar Anaebaena-Azolla symbiosis "green manure" Inoculation of 0.2 kg of Azolla per hectare is equivalent to 30 kg/hectare of commercial nitrogen fertilizer. Not as popular now because we have found that Azolla growth is limited by phosphate, which is non-renewable. Next break is low-phosphorus requiring Azolla Ascophyllum nodosum (bladder wrack) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Calcareous algae to reduce soil acidity. Caulerpa racemosa (Nama) in Fiji food Chlamydomonas and other soil algae soil amendment by making mucilage that loosen compacted soil. Chlorella only microscopic green used commercially Chondrus crispus (Irish Moss) agar, carrageenan Codium geppii (Sagati) in Fiji food Diatom fossils (diatomaceous earth, diatomite) flea killers, embalming compounds, enamel polishes, diatomaceous bricks, to filter sewage in UK from 1976, to filter sugar cane liquors from 1914; Nobel stabilized nitroglycerine. Durvillaea (Australia & Chile) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Ecklonia alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Eucheuma in Philippines carrageenan Furcillaria fastigiata in Denmark agar Gelidium amansii in Japan agar Gelidium arborescens in USA agar Gelidium cartillagineum in USA agar Gelidium latifolium in Ireland. agar Gelidium nudifrons in USA. agar Gelidium pulchellum Ireland agar Gigartina clavifera in New Zealand carrageenan Gigartina mamillosa (very similar and known as Irish Moss). carrageenan Gigartina undulata in New Zealand carrageenan Gracilaria (Lumicevata) in Fiji food Hypnea (Lumiwawa) in Fiji food Kelp and mixed seaweeds liquid fertilizers, soil amendment Wayne and Wanda! rich in potassium and nitrogen but low in phosphate. Free of terrestrial weeds and fungi. Kelps (Kombu) vegetables in Japan and China Laminaria (kelp - English; kombu - Japan and China) Food. Maybe 75% of their dry bulk is indigestible Laminaria digitata in UK and Norway alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Laminaria cloustoni (oar weed) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Macrocystis pyrifera (kelp - USA) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Mastocarpus stellata in British Isles. agar Phyllophora nervosa in USSR. agar Porphyra (laver - England and USA; luche - Chile; karengo - Maori; nori - Japan; slack - Scotland; sloke - Ireland) Food - Maybe 75% of their dry bulk is indigestible. Toasted and used to wrap sushi North Pacific. Cultivated for centuries in Japan, Korea and China Pterocladia agar Spirulina food protein - blue-green used for protein food in old (Lake Chad) and new world (Aztecs before conquistadores) Ulva food - eaten as a green vegetable Undaria pinnatifida (wakame) food - Maybe 75% of their dry bulk is indigestible http://seaweed.ucg.ie/Algae/fucus.html Fucus vesiculosis

Or. Fucales Southern Hemisphere Wrack or Rockweed 4 Families Fucaceae (Fucus, Ascophyllum) Sargassaceae (Sargassum, Turbinaria) Cystoseiraceae Hormosiraceae (Hormosira) Turbinaria ornata *Oceanographic conditions determine these 3 factors Next to coral reefs, considered largest biological structure in the oceans Useful species, common names, some of their uses? SpeciesCommon nameUses acanthopeltis agar Ahnfeltia plicata in USSR agar Anaebaena-Azolla symbiosis "green manure" Inoculation of 0.2 kg of Azolla per hectare is equivalent to 30 kg/hectare of commercial nitrogen fertilizer. Not as popular now because we have found that Azolla growth is limited by phosphate, which is non-renewable. Next break is low-phosphorus requiring Azolla Ascophyllum nodosum (bladder wrack) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Calcareous algae to reduce soil acidity. Caulerpa racemosa (Nama) in Fiji food Chlamydomonas and other soil algae soil amendment by making mucilage that loosen compacted soil. Chlorella only microscopic green used commercially Chondrus crispus (Irish Moss) agar, carrageenan Codium geppii (Sagati) in Fiji food Diatom fossils (diatomaceous earth, diatomite) flea killers, embalming compounds, enamel polishes, diatomaceous bricks, to filter sewage in UK from 1976, to filter sugar cane liquors from 1914; Nobel stabilized nitroglycerine. Durvillaea (Australia & Chile) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Ecklonia alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Eucheuma in Philippines carrageenan Furcillaria fastigiata in Denmark agar Gelidium amansii in Japan agar Gelidium arborescens in USA agar Gelidium cartillagineum in USA agar Gelidium latifolium in Ireland. agar Gelidium nudifrons in USA. agar Gelidium pulchellum Ireland agar Gigartina clavifera in New Zealand carrageenan Gigartina mamillosa (very similar and known as Irish Moss). carrageenan Gigartina undulata in New Zealand carrageenan Gracilaria (Lumicevata) in Fiji food Hypnea (Lumiwawa) in Fiji food Kelp and mixed seaweeds liquid fertilizers, soil amendment Wayne and Wanda! rich in potassium and nitrogen but low in phosphate. Free of terrestrial weeds and fungi. Kelps (Kombu) vegetables in Japan and China Laminaria (kelp - English; kombu - Japan and China) Food. Maybe 75% of their dry bulk is indigestible Laminaria digitata in UK and Norway alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Laminaria cloustoni (oar weed) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Macrocystis pyrifera (kelp - USA) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Mastocarpus stellata in British Isles. agar Phyllophora nervosa in USSR. agar Porphyra (laver - England and USA; luche - Chile; karengo - Maori; nori - Japan; slack - Scotland; sloke - Ireland) Food - Maybe 75% of their dry bulk is indigestible. Toasted and used to wrap sushi North Pacific. Cultivated for centuries in Japan, Korea and China Pterocladia agar Spirulina food protein - blue-green used for protein food in old (Lake Chad) and new world (Aztecs before conquistadores) Ulva food - eaten as a green vegetable Undaria pinnatifida (wakame) food - Maybe 75% of their dry bulk is indigestible Sargassum

Or. Durvilleales Durvillea antarctica with lesions – Southern hemisphere only

MS Phaeophyta (1957) Ectocarpales – Ectocarpus (6sp) Sphacelariales - Sphacelaria Dictyotales – Dictyota (2), Padina Fucales – Sargassum (3) about 13 species…

Rhodophyte taxonomy Two subclasses: Bangiophycidae and Florideophycidae (Dawes 1998). Bangiophycidae Uninucleate cells Single stellate, central plastid Intercalary (diffuse) cell division Absence of pit connections Mostly asexual reproduction Simple unicell – multicell thallus forms 3 Orders Florideophycidae multinucleate cells in many species Several to many discoid chloroplasts per cell Cell division is atypical Presence of pit plugs Sexual reproduction common Only multicell thalli 9 Orders

Rhodophyte taxonomy Two subclasses: Bangiophycidae and Florideophycidae (Dawes 1998). Bangiophycidae Uninucleate cells Single stellate, central plastid Intercalary (diffuse) cell division Absence of pit connections Mostly asexual reproduction Simple unicell – multicell thallus forms 3 Orders Florideophycidae multinucleate cells in many species Several to many discoid chloroplasts per cell Cell division is atypical Presence of pit plugs Sexual reproduction common Only multicell thalli 9 Orders

THALLUS COARSLEY - BRANCHED FILAMENT SHEET ENCRUSTING JOINTED - CALCAREOUS THICK - LEATHERY

Thallus organization Unicellular - Porphyridium Bangiophycidae Unicellular - Porphyridium Filamentous – Erythrocladia, Polysiphonia Sheet-like – Porphyra Branching – Laurencia, Gracilaria Encrusting calcareous – Lithothamnion, Melobesia Jointed Calcareous – Amphiroa, Corallina Taxonomy based on morphology, reproductive structures, and life cycles

Macroalgae - Rhodophyta

Macroalgae - Rhodophyta

Subclass Florideophycidae taxonomy Dawes, pg 153

Or. Palmariales Dulse Palmaria palmata

Or. Nemaliales – 4 Families Galaxaura Liagora Scinaia

Or. Gelidiales – 2 Families Gelidium Gelidiella

Or. Bonnemaisoniales Asparagopsis - gametophyte Falkenbergia - sporophyte

Or. Cryptonemiales – 12 Families Grateloupia Cryptonemia Halymenia

Or. Corallinales – 1 Family w 35 genera Articulated (geniculate) Non-Articulated (encrusting) Porolithon Neogoniolithon Melobesia Jania Corallina Amphiroa Lithothamnion

Or. Gigartinales – 28 Families Hypnea Eucheuma Kallymenia Gracilaria

Or. Rhodymeniales – 3 Families Botryocladia Champia Chrysymenia

Or. Ceramiales – 4 Families Ceramiaceae (100 genera) Callithamnion

More Fam. Ceramiaceae Ceramium

More Fam. Ceramiaceae Ceramium Ceramium

More Fam. Ceramiaceae Ceramium Centroceras

Or. Ceramiales – 4 Families Ceramiaceae (100 genera) Callithamnion Delesseiraceae (90 genera) Caloglossa Dasyaceae (10 genera) Dasya Rhodomelaceae (125 genera) - Laurencia

Or. Ceramiales – 4 Families Ceramiaceae (100 genera) Callithamnion Delesseiraceae (90 genera) Caloglossa Dasyaceae (10 genera) Dasya Rhodomelaceae (125 genera) - Laurencia Polysiphonous construction

Polysiphonia (Fam. Rhodomelac)

Macroalgae - Rhodophyta

MS Rhodophyta (1957) Bangiales – Erythrotrichia, Goniotrichum Nemalionales – Achrochaetium (2sp) Gelidiales – Gelidium (2) Cryptonemiales - Grateloupia Corallinales – Fosliella (2) Gigartinales – Gymnogongrus, Agardhiella, Hypnea, Gracilaria Ceramiales – Ceramium, Spyridia, Caloglossa, Bostrichia (2), Chondria, Herposiphonia, Polysiphonia (3), Lophosiphonia about 24 species…

AlgaeBase.org

Seaweeds in your diet!

Cultivation of Red Algae – carragenans, agar Eucheuma cultivation – Zanzibar, E. Africa

Kelps for algin (thickener) http://www.starthrower.org/research/kelpmisc/kelp_mp.htm Kelps for algin (thickener) Used in over 300 products: ice cream, paints, sauces, and toothpaste California/Oregon – heavily regulated 8m/20ft wide swaths 550 metric tons/day/ship 700’000 metric tonnes p.a. in China alone!

http://life.bio.sunysb.edu/marinebio/kelpforest.html Kelps Coral Reefs

KELP FORESTS

What Charles Darwin said… “I can only compare these great aquatic forests…with the terrestrial ones in the intertropical region. Yet if in any country a forest was destroyed, I do not believe nearly so many species of animals would perish as would here, from the destruction of the kelp. Amidst the leaves of this plant numerous species of fish live, which nowhere else could find food or shelter…” Charles Darwin, 1 June 1834, Tierra del Fuego, Chile

Kelp forests vs terrestrial forests Kelp forest structure similar to terrestrial forests – different canopy levels, varied understory Kelp forests more productive, more diverse (phylogenetically) Shorter life spans, shorter heights Kelp forests reach 10-30 meters within 1-3 years, few sp. lasting more than 25 years Terr. forests reach 10-30 meters within 20-30 years, living thousands of years Recruitment and growth of new kelp depends upon canopy breaks for available light Terrestrial forests are habitat for roughly 3 phyla Kelp forests are habitat for 10 or more phyla

Similarities and differences to terrestrial forests Both forest types: Recruitment and growth depends upon canopy breaks for available light Structure similar - different canopy levels, varied understory Form foundation for large diversity of animals and plants Both are primary producers, get their energy from the sun Kelp forests have shorter life spans, shorter heights Kelp forests are faster growing, shorter-lived: Few kelp sp. last more than 25 years Terr. forest trees can live for thousands of years Kelp forests are more diverse in terms of number of animal phyla, less in terms of animal species? Terrestrial forests are habitat for roughly 3 phyla Kelp forests are habitat for 10 or more phyla Kelp forest structure similar to terrestrial forests – different canopy levels, varied understory Kelp forests more productive, more diverse (phylogenetically) Shorter life spans, shorter heights Kelp forests reach 10-30 meters within 1-3 years, few sp. lasting more than 25 years Terr. forests reach 10-30 meters within 20-30 years, living thousands of years Recruitment and growth of new kelp depends upon canopy breaks for available light Terrestrial forests are habitat for roughly 3 phyla Kelp forests are habitat for 10 or more phyla

Kelp: Division Phaeophyta (Brown Algae) Brown Algae (Phaeophyta) 1000 species, almost all marine Includes Sargassum, Padina, kelps, rockweeds Most common in cold, temperate seas Two pigments for photosynthesis: 1) Chlorophyll a (like all plants) 2) Fucoxanthin (brown color)

Kelp: Division Phaeophyta (Brown Algae) Parts of a kelp Leaflike blades Gas-filled pneumatocysts Long hollow stem or stipe Rootlike holdfast to attach to substrate Complex life cycle *Oceanographic conditions determine these 3 factors Next to coral reefs, considered largest biological structure in the oceans Useful species, common names, some of their uses? SpeciesCommon nameUses acanthopeltis agar Ahnfeltia plicata in USSR agar Anaebaena-Azolla symbiosis "green manure" Inoculation of 0.2 kg of Azolla per hectare is equivalent to 30 kg/hectare of commercial nitrogen fertilizer. Not as popular now because we have found that Azolla growth is limited by phosphate, which is non-renewable. Next break is low-phosphorus requiring Azolla Ascophyllum nodosum (bladder wrack) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Calcareous algae to reduce soil acidity. Caulerpa racemosa (Nama) in Fiji food Chlamydomonas and other soil algae soil amendment by making mucilage that loosen compacted soil. Chlorella only microscopic green used commercially Chondrus crispus (Irish Moss) agar, carrageenan Codium geppii (Sagati) in Fiji food Diatom fossils (diatomaceous earth, diatomite) flea killers, embalming compounds, enamel polishes, diatomaceous bricks, to filter sewage in UK from 1976, to filter sugar cane liquors from 1914; Nobel stabilized nitroglycerine. Durvillaea (Australia & Chile) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Ecklonia alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Eucheuma in Philippines carrageenan Furcillaria fastigiata in Denmark agar Gelidium amansii in Japan agar Gelidium arborescens in USA agar Gelidium cartillagineum in USA agar Gelidium latifolium in Ireland. agar Gelidium nudifrons in USA. agar Gelidium pulchellum Ireland agar Gigartina clavifera in New Zealand carrageenan Gigartina mamillosa (very similar and known as Irish Moss). carrageenan Gigartina undulata in New Zealand carrageenan Gracilaria (Lumicevata) in Fiji food Hypnea (Lumiwawa) in Fiji food Kelp and mixed seaweeds liquid fertilizers, soil amendment Wayne and Wanda! rich in potassium and nitrogen but low in phosphate. Free of terrestrial weeds and fungi. Kelps (Kombu) vegetables in Japan and China Laminaria (kelp - English; kombu - Japan and China) Food. Maybe 75% of their dry bulk is indigestible Laminaria digitata in UK and Norway alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Laminaria cloustoni (oar weed) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Macrocystis pyrifera (kelp - USA) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Mastocarpus stellata in British Isles. agar Phyllophora nervosa in USSR. agar Porphyra (laver - England and USA; luche - Chile; karengo - Maori; nori - Japan; slack - Scotland; sloke - Ireland) Food - Maybe 75% of their dry bulk is indigestible. Toasted and used to wrap sushi North Pacific. Cultivated for centuries in Japan, Korea and China Pterocladia agar Spirulina food protein - blue-green used for protein food in old (Lake Chad) and new world (Aztecs before conquistadores) Ulva food - eaten as a green vegetable Undaria pinnatifida (wakame) food - Maybe 75% of their dry bulk is indigestible

Kelp “life history” Kelp alternates between a large sporophyte and a tiny gametophye - sporophyte = “plant that makes spores” (diploid, 2N, large plant-like stage) - gametophyte = “plant that makes gametes” (haploid, 1N, small inconspicous stage - can be a tiny thread) http://www.mbari.org/staff/conn/botany/browns/james/Web/lifehis.htm *Oceanographic conditions determine these 3 factors Next to coral reefs, considered largest biological structure in the oceans Useful species, common names, some of their uses? SpeciesCommon nameUses acanthopeltis agar Ahnfeltia plicata in USSR agar Anaebaena-Azolla symbiosis "green manure" Inoculation of 0.2 kg of Azolla per hectare is equivalent to 30 kg/hectare of commercial nitrogen fertilizer. Not as popular now because we have found that Azolla growth is limited by phosphate, which is non-renewable. Next break is low-phosphorus requiring Azolla Ascophyllum nodosum (bladder wrack) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Calcareous algae to reduce soil acidity. Caulerpa racemosa (Nama) in Fiji food Chlamydomonas and other soil algae soil amendment by making mucilage that loosen compacted soil. Chlorella only microscopic green used commercially Chondrus crispus (Irish Moss) agar, carrageenan Codium geppii (Sagati) in Fiji food Diatom fossils (diatomaceous earth, diatomite) flea killers, embalming compounds, enamel polishes, diatomaceous bricks, to filter sewage in UK from 1976, to filter sugar cane liquors from 1914; Nobel stabilized nitroglycerine. Durvillaea (Australia & Chile) alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Ecklonia alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Eucheuma in Philippines carrageenan Furcillaria fastigiata in Denmark agar Gelidium amansii in Japan agar Gelidium arborescens in USA agar Gelidium cartillagineum in USA agar Gelidium latifolium in Ireland. agar Gelidium nudifrons in USA. agar Gelidium pulchellum Ireland agar Gigartina clavifera in New Zealand carrageenan Gigartina mamillosa (very similar and known as Irish Moss). carrageenan Gigartina undulata in New Zealand carrageenan Gracilaria (Lumicevata) in Fiji food Hypnea (Lumiwawa) in Fiji food Kelp and mixed seaweeds liquid fertilizers, soil amendment Wayne and Wanda! rich in potassium and nitrogen but low in phosphate. Free of terrestrial weeds and fungi. Kelps (Kombu) vegetables in Japan and China Laminaria (kelp - English; kombu - Japan and China) Food. Maybe 75% of their dry bulk is indigestible Laminaria digitata in UK and Norway alginates thickening agents (fruit drinks and salad dressing), colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Laminaria cloustoni (oar weed) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Macrocystis pyrifera (kelp - USA) alginates thickening agents (fruit drinks and salad dressing) colloid stabilizers in food (ice creams, sherbets and cheeses), textile, cosmetic, pharmaceutical, paper and welding industries. Mastocarpus stellata in British Isles. agar Phyllophora nervosa in USSR. agar Porphyra (laver - England and USA; luche - Chile; karengo - Maori; nori - Japan; slack - Scotland; sloke - Ireland) Food - Maybe 75% of their dry bulk is indigestible. Toasted and used to wrap sushi North Pacific. Cultivated for centuries in Japan, Korea and China Pterocladia agar Spirulina food protein - blue-green used for protein food in old (Lake Chad) and new world (Aztecs before conquistadores) Ulva food - eaten as a green vegetable Undaria pinnatifida (wakame) food - Maybe 75% of their dry bulk is indigestible

Kelp forest zonation

Kelp forests occur in cool water

Kelp forests - Where do they occur? Cold, temperate water Kelp forests are more common off West Coast of N. America than E. Coast Why? because of cool California current off Ca coast vs.warm Gulf Stream on East Coast

The ecological role of kelp Dampen wave action Reduce coastal erosion Enhance recruitment of fish and invertebrates High productivity and turnover of plant matter puts many nutrients into system High structural diversity (like a forest on land) provides shelter and habitat for many species Habitat complexity created: - understory with filtered light - canopy at surface Kelp forests alter and shape local environments and ecologies -- dampen wave movement, influencing water flow, coastal erosion, sedimentation, benthic productivity (primary & secondary), & recruitment of species -- canopy coverage reduces light, creating conditions for low light intensity species of algae and animal Sessile algae and animals use kelp as substrate Mobile animals live and feed off of kelp – some cannot live without kelp Predatory fishes use kelp forests as primary habitat Canopy litter acts as food for detritivores Predatory fishes use kelp forests as primary habitat – canopy loss can result in trophic cascades Canopy litter acts as food for detritivores on sea floor – direct contribution to secondary productivity

Kelp forests come and go… Kelp forests can disappear quickly -- temperature changes -- severe storms -- outbreaks of herbivorous invertebrates -- disease However, the kelp forest community can regrow just as quickly

Kelp forests are deforested by natural causes Salinity, temperature change, storms, seasons Can recover fairly quickly under natural conditions Oldest known term for algal deforestation, means “rock-burning” -- 1902 -- Japanese coast, reason: salinity anomalies -- lower latitude deaths caused by environmental changes -- destruction is short-lived, reversible Reasons for death are different depending on latitude placement LOWER LATITUDE DEATH (less than 40 degrees): deforestations result from temperature, salinity and nutrient anomalies which directly kill or trigger diseases MID-LATITUDE DEATH (40-60 degrees): kelp deforestation caused by intense herbivory by sea urchins

Sea urchins graze on kelp Primary cause for mid-lat. Death “Urchin barrens” – 1960s Urchin-caused deforestation most often seen in Northern Hemisphere (particularly Alaska) In Southerm Hem., widespread urchin death prevented by increased competition, oceanographic limitations, increased predation

Sea otters control urchin populations by eating them Kelp forest habitat determined by urchin abundance Urchin abundance determined by various factors: Predation (ex. Otters, fish -- most important factor), interspecific competition (other herbivores, i.e. abalone), disease, turbulence, storms, etc.

The last of the otters 150 years after excessive hunting, by late 1800s, only 3000 otters left International Fur Treaty established in 1911: illegal to kill sea otters, even for indigenous Alaskans Sea otter population drops from previous 300000 to 3 thousand by late 1800s – only found in Alaska, gone from rest of range People start to realize that the otter may go extinct after nearly 150 years of excessive hunting and decide to protect it 1911 – International Fur Treaty is established, making it illegal to kill sea otters, even for Alaskan indigenous people a. Distribution of sea otters before fur harvest began in 1741 and populations that survived the harvest, providing the nucleus for recovery of the species. Width of shaded area is not relative to sea otter habitat.

Otters repopulate In 1980s, census conducted: 150,000 otters on earth, mostly in Alaska 2000 otters found in California, population was thought to be extinct there Although sea otters have slow gestation periods (6 months) and only usually give birth to 1 infant (twins are exceptions), pop. slowly builds up By 1980’s, pop. assessed: nearly 150000 otters alive on Earth, primarily in Alaska Pop. of 2000 discovered in California after thought to be locally extinct Worldwide population thought to be at safe equilibrium density Sea otters give birth to 1 infant per pregnancy, 6 month gestation

Kelp: alternate stable foodwebs “Keystone spp”

A new threat emerges In 1991, first killer whale attack on Alaskan otter is witnessed In 1991, the first attack by a killer whale on a sea otter is witnessed by otter ecologists – not considered significant, just surprising In 1994, population analysis of Alaskan Islands reveals a drop of 90% in several locations By 1998, Alaskan otter population estimated to have dropped dramatically to 45,000 from previous 148,000 Scientists struggle to find answer

Sequential overharvesting of marine mammals Killer whales may have switched to otters because their normal food (whales and seals) has become rare. The chance that a change in fishing practices could eventually result in replenished stocks of sea lion and fur seal fish prey, resulting in change back of killer whales to natural prey, and sea otters grow again Fishing practices will not change Chances of killer whales changing back, anyway, are not high Time it takes for fish pop. to rebuild and chance for fur seals and sea lions to recover will be too long and might not be possible

Kelp foodwebs and Alternate Stable States (A.S.S.) Killer whales

A. S. S. - Kelps Kelp dominated, many otters Large disturbance, “Unatural”, entire food wed has changed. Small disturbance, “Natural” Kelp dominated, many otters Nutrient poor, few urchins Kelp dominated, few otters Nutrient poor, few urchins Algal dominated, many otters Nutrients elevated, lots urchins Algal dominated, few otters, Nutrient rich, lots urchins

A. S. S. – Coral Reefs Urchin die-off & Over-fishing route Algal dominated, some fish Nutrient poor, some urchins Coral dominated, many fish Nutrient poor, many urchins Eutrophication & Over fishing route Algal dominated, few fish Nutrient rich, few urchins Calc Red Algae dominated, some fish Nutrient rich, many urchins

References Dayton, PK, Tegner MJ, Edwards PB, et al Sliding baselines, ghosts, and reduced expectations in kelp forest communities ECOL APPL 8 (2): 309-322 MAY 1998 Estes, JA, Duggins, DO, Rathbun, GB. The ecology of extinctions in kelp forest communities. CONSERV BIOL 3 (3): 252-264 SEP 1989 Jackson, JBC, Sala E Unnatural oceans SCI MAR 65: 273-281 Suppl. 2 SEP 2001 Jackson, JBC, Kirby, MX, Berger, WH, et al. Historical overfishing and the recent collapse of coastal ecosystems SCIENCE 293 (5530): 629-638 JUL 27 2001 Jackson, JBC What was natural in the coastal oceans? P NATL ACAD SCI USA 98 (10): 5411-5418 MAY 8 2001 Simenstad CA, Estes JA, Kenyon KW. Aleuts, sea Otters, and alternate stable-state communities SCIENCE 200 (4340): 403-411 1978 Steneck RS, Graham MH, Bourque BJ, et al. Kelp forest ecosystems: biodiversity, stability, resilience and future ENVIRON CONSERV 29 (4): 436-459 DEC 2002 Tegner, MJ, Dayton, PK Sea-urchins, El-Ninos, and the long-term stability of southern California kelp forest communities MAR ECOL-PROG SER 77 (1): 49-63 OCT 1991

Summary Kelps largest protists - to 70m long! Kelp habitats – Brown canopy, Red understory Browns taxonomy by reproduction and life-history Reds taxonomy by morphology, reproduction, and life-history Human uses of Kelps and Red algae for food additives.