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Kelp Bed / I. What are beds or ? Stands of large brown in the order Laminariales I. What are kelp beds and forests? (i.e., ) that provide food and shelter for a diverse II. Distribution array of marine and animals. III. Morphology IV. Kelp life cycle V. Biological characteristics of kelp forests VI. Abiotic Influences VII. Effects of kelp on biotic conditions VIII.Biotic interactions IX. Kelp harvesting

Kelp Forest vs. Kelp Bed I. What are kelp beds/forests? -- terms often used interchangeably, but technically: 27 genera of kelps form kelp beds or forests •kelp forest = large that form floating surface canopies

•kelp bed = large brown algae that do not form a surface canopy

kelp forest kelp bed Pelagophycus

Pterygophora

II. Distribution II. Distribution • worldwide in temperate (cold & nutrient rich) North and South American west coasts: – larger forests in water <20˚ C • kelp forests are dominated by two genera of canopy-forming kelps: • generally shallower than 15-40 m – (bull kelp) in north – Macrocystis (giant kelp) in north & south

Macrocystis Nereocystis

1 II. Distribution II. Distribution

• on the North American east coast, Asian east coast, and • in , , and , kelp in Europe, kelp beds are dominated by beds are dominated by Ecklonia

III. Kelp Morphology IV. Kelp Life Cycle spores

• holdfast - rootlike attachment to Alternation of generations, substrate haploid and diploid:

• stipe – long stem gametes • diploid sporophytes fertilization • blades - “leaves” produce haploid spores • - gas-filled floats • haploid gametophytes (male & female) produce haploid gametes that fuse to form a diploid sporophyte

V. Biological Characteristics of Kelp Forests V. Biological Characteristics of Kelp Forests: • High Productivity - Macrocystis can grow up to 50 cm per day & reaches 60 m long Comparisons between terrestrial forests and giant kelp forests FOREST TYPE NET PRODUCTION PRODUCER MASS LITTER MASS (dry kg / m2 / yr) (dry kg / m2 ) (dry kg / m2 ) • High diversity Tropical rain forest 2.2 45 0.2

Tropical deciduous forest 1.6 35 0.5

Temperate evergreen forest 1.3 35 3.0

Temperate deciduous forest 1.2 30 2.0

Boreal forest 0.8 20 4.0

Giant kelp forest 2.2 0.35 0.015

2 V. Biological Characteristics of Kelp Forests: Percentage of Kelp-Derived Carbon in Kelp Forest Consumers (from Duggins et al. 1989, Alaskan kelp forests, & Laminaria) Kelp as food: 13C to 12C ratios • kelps are ~ 3-10x more productive than shallow-water Note: more recent research indicates these estimates are too & about 50x more productive than oceanic phytoplankton high because of problems with the original methods (Miller and Page 2012) • but few animals directly eat the living fronds of kelps Suspension Feeders % kelp carbon • as kelp disintegrates, chunks, particles, and organic molecules feed mussel (Mytilus edulis) 25 - 40 suspension feeders and deposit feeders soft coral (Alcyonaria sp). 40 - 70 barnacle (Balanus nubilus) 75 - 85 anemone (Metridium senile) 15 - 40 rock jingle (Pododesmus cepio) 40 - 55 mysid (Proneomysis sp.) 45 - 60 amphipod (Anonyx sp.) 70 - 95 (Dermaturus mandtii) 20 - 35 Predators rock greenling (Hexagrammos lagocephalus)) 40 - 65 sea star (Leptasterias spp.) 30 - 55 cormorant (Phalacorcorax peligicus) 35 - 70

And kelp forests/beds Export their Production to other : V. Biological Characteristics of Kelp Forests: e.g.,Kelp Wrack Diversity

High Diversity of Primary Producers – not just kelps

V. Biological Characteristics of Kelp Forests: Causes of High Diversity Diversity • high structural diversity (3-dimensional High Diversity of Consumers structure; like coral reefs and tropical rain forests) – places to hide from predators – unique locations

• high productivity

3 Microhabitats & associated species in a typical California kelp Canopy assemblage: forest: isopods, tubeworms, bryozoans, A. Canopy Membranipora membranacea,  filter feeding colonies; eaten by several species assemblage a common bryozoan of and a few fishes

B. Planktonic assemblage

Idotea sp., a common isopod C. Holdfast assemblage

D. Horizontal substrate assemblage

E. Vertical wall assemblage

Canopy assemblage: fishes Holdfast assemblage: sea urchins, crustaceans, worms, brittle stars,

kelp bass giant kelpfish

kelp perch

brittlestars in señorita blue rockfish holdfast

Horizontal substrate assemblage: Vertical wall assemblage: urchins, sea stars, algae, and bottom coralline algae, anemones, cup corals, sponges, mollusks in crevices

blackeye goby

sculpin

4 VI. Abiotic factors that influence kelp forests VI. Abiotic factors that influence kelp forests

• Light • Temperature LIGHT • Nutrients • Water Motion • Hard Substrate

availability of light limits depths of kelps: • need > 1% of surface irradiance • can grow deeper in clearer water

VI. Abiotic factors that influence kelp forests VI. Abiotic factors that influence kelp forests

Water Motion: Substrate type: • big waves rip out kelp forests • kelps generally require a hard substrate to attach to (rocks) • some species better able to deal • cannot settle and grow on sand or other soft substrates with high water motion than others: - e.g., Nereocystis can handle larger waves (water motion) than Macrocystis

VI. Abiotic factors that influence kelp forests Temperature & Nutrients: Effects of El Niño low temperature Temperature & Nutrients: high nutrients El Niño conditions: • correlated: high temperature  low nutrients • warm surface waters • nitrate thought to be key limiting nutrient • depress thermocline • keep nutrients away from kelp high temperature Kelp growth declines & then kelps die low nutrients

from Dayton et al. 1999

5 Variation in abiotic conditions often drives kelp Seasonal variation • example: Macrocystis in Southern California -highest abundance & during winter/spring (cold water, high nutrients) • Seasonal variation (predictable) -lowest abundance late summer/fall (warm water, low nutrients) • Interannual variation (unpredictable)

Dayton et al. 1992

Interannual variation Interannual variation • example: Macrocystis in Southern California (Dayton et al. 1992)

- highest abundance during cold water periods 1948 (La Niña) Pt. Loma Kelp Forest: - lowest abundance during warm water periods 1940’s - 1970’s (El Niño)

1959

1965

1970

VIII. Biological factors that influence kelp forests VII. Effects of kelp beds on abiotic conditions GRAZING

• kelp forests alter the local hydrodynamic regime — slow down currents — allow sediments to settle out — “filter” edible particles (e.g., )

6 Competition: Trophic Interactions

• intense competition for light - occurs among kelps & understory algae X - in southern CA, Macrocystis is the dominant competitor because of fast growth

• competition for space - competition for attachment spots on hard substrate - occurs among algae and also between algae and

Alternate Stable States? changes kelp forests and urchin barrens

X

X

Factors that influence populations in IX. Kelp Harvesting southern California

• kelp is a main source of algin and other emulsifiers and • other predators on sea urchins in California thickeners - foods - - cosmetics - - paints - pharmecueticals • sporadic recruitment of sea urchins in California • $40 million a year industry in California in the mid 1980’s • urchin disease - known to greatly reduce urchin numbers over relatively wide • started in early 1900’s to make potash for gunpowder in areas World War I - in California these outbreaks of disease are generally associated with periods of warm water - expansion of kelp beds have been documented following massive urchin die offs

7 IX. Kelp Harvesting Summary

• boats with cutters about six feet deep take all surface growth Kelp forests are… and the top 4 feet of the stipes • highly productive • kelp beds were leased to private groups • highly diverse • in California, Kelco (until 2005) • highly dynamic • between 100,000 and 170,000 - due to strong biotic & abiotic influences wet tons were harvested from California waters each year • no longer harvested in CA

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