Life in the rocky subtidal, rocky reefs, kelp forests

Chapter 15 (pgs. 418-432)

Jake Schiferl and Jessie Cartwright

Introduction: What are Rocky Reefs and Kelp Forests?
Rocky Reef Habitat: ‡ Rocks occur above or below the water line ‡ As the rocks erode, cracks and holes appear that create shelter for living things o Different rock types weather away in different ways and, therefore, support different species ‡ Sometimes, parts of the reef are exposed at low tide (see discussion of rocky intertidal) o Can create rock pools o Some species can survive out of the water during low tide ‡ Subtidal is generally submerged, usually only exposed during extreme low tides ‡ Kelp forests are generally formed below the tide level of a rocky reef

Rocky reefs and kelp forests are generally interlinked!!!

Introduction: Importance of the Topic
‡ Rocky reefs and kelp forests support many species o Biodiversity "hotspotsµ o Hotspot- a threatened region with a large range of endemic (species native or restricted to a certain area) plant species (at least 1,500) but where more than 70% of the original habitat has been lost due to human activities. Typically, diversity of endemic vertebrates is also high. o Jeffrey S. Levinton- discusses hotspots in The Hudson River Estuary o Ch. 20: ´Scientists, managers, and educators are interested in estuarine wetlands because they are ¶hotspots· of ecological processes, biological diversity, and human activity and impact within the estuary.µ o Includes many fisheries species o Good for tourism ‡ Over-exploitation and introduced invasives (human impacts) o Loss of biodiversity [including fisheries species (impact on humans)]

Introduction: Importance of the Topic
‡Healthy rocky reef/kelp forest ecosystems (left) are very productive and support a great deal of biodiversity. ‡However, each part of the ecosystem is interconnected so if one part is negatively affected, this can also have negative impacts on the rest of the ecosystem ‡Example: decrease in sea otter populations due to fur hunting leads to increased sea urchin populations (sea otter prey) which leads to decreased giant kelp (what sea urchins feed on) ‡Additionally, overharvesting aids in the reduction of kelp populations [eventually, barrens (bare bottoms) are formed] ****MORE ON THIS TOPIC LATER!****

Introduction: What are Rocky Reefs and Kelp Forests and why are they important? Overview of the kelp forest .

g.) Graph from textbook (pg. grazers (urchins. sea squirts.). etc.The Rocky Reef-Kelp Forest System ‡ Dominated by kelps and seaweeds in shallower waters o Due to light limitations o Foundation species that alter water flow and continuously provide detrital particles used by many invertebrates ‡ Dominated by epifaunal/sessile animals in deeper waters o Epifaunal: an animal living on the surface of the seabed. anemones. etc. ‡ Mobile organisms dominate in the transition between shallower and deeper waters o Predatory fish. snails. abalones. 419): ‡ kelp and algae dominate lower depths ‡ animals such as invertebrates dominate higher depths . invertebrate predators (sea stars. bryozoans. etc. generally attached to submerged animals or plants on the hard substrate (sessile)  e. nudibranchs.

sessile organisms .The Rocky Reef-Kelp Forest System A fun example: ‡ Shallow Waters: kelps and seaweeds ‡ Transition: mobile organisms (fish and other predators) ‡ Deeper Waters: epifaunal.

Subtidal Rocky Reefs ‡ Mainly in temperate and higher-latitude environments o Subtidal rocky outcrops occur in the tropics but are usually dominated by corals and other large calcareous organisms ‡ Subtidal rocky reefs tend to be dominated by colonial (modular)animals o o Examples: colonial bryozoans. hydroids. sponges. bivalves. and sea squirts. as well as. and barnacles Intertidal rocky shores dominated by individuals such as barnacles and mussels. individuals such as anemones. .

barnacles. mussels can readily smother other species) Colonial anemone anemone outbreak anemone polyps ..g. and bivalves) result in death ‡ Leading to modular species dominance ‡ Not self evident that animals that occur as individuals are inherently competitively inferior (e. or zooids of a bryozoan ‡ Ability to spread along a surface helps to prevent predation and competition o Predation on colonial animals will result in a recolonization of the area o Predation on animals that occur as individuals (e.g. anemones..g. mussels. polyps of a colonial anemone.Subtidal Rocky Reefs ‡ Modular organisms (textbook definition): consist of interconnected and usually identical units o e.

so kelp and colonial animals mostly limited to subtidal ‡ Water below the surface layer the temperature is predictably cool and salinity is fairly constant year around .Subtidal Rocky Reefs ‡ Physiological limitations on kelp and colonial animals to subtidal rocky reefs o Not capable of closing off soft tissue from extreme temperature and salinity changes o Desiccation in the intertidal.

Subtidal Rocky Reefs Structure ‡ Rocky subtidal fauna are partly limited to subtidal depths due to: o Inability to survive the low salinity o Strong pycnocline presents a barrier for dispersal of larvae  Pynocline (textbook definition): Depth zone within which seawater density changes pynocline example .

Subtidal Rocky Reefs ‡ Rocky subtidal reefs are very patchy.g. with two main patch types on rocks o On vertical walls. patch size limited by interspecific and intraspecific competition  Patches with rich assemblage of sessile marine invertebrates o On horizontal walls. sedimentation)  Patch dominated by turf-forming red calcareous algae (resistant to predation) or green algae (in shallower areas)  Very few sessile invertebrates red calcareous algae Ulva (green algae) . landslides. patch size limited by biological or physical disturbance (e..

bryozoan larvae.Subtidal Rocky Reefs ‡ Case Study of Patchiness in subtidal rocky communities: Franz Smith and Jon Witman (1999) o Placed settling plates within and at various distances from the different patches o Found settlement to be much higher within the patches and decreased as distance from the patch increased  Settlement of short-lived larvae such as ascidian tadpole larvae. and hydroid actinula larvae o Discovered that larval recruitment from the local patch itself was sufficient to perpetuate the local patches  Many larvae are lecithotrophic  Lecithotrophic larvae (textbook definition): a planktonic-dispersing larva that lives off yolk supplied via the egg Lecithotrophic development Lecithotrophic Life Cycle settling plate .

Subtidal Rocky Reefs Competitive Interactions ‡ Competitive success: ability to overgrow neighbors and to resist overgrowth by other species ‡ Study of a species of ascidian. in general. o Rapidly growing species were good at overgrowing but were not good at resisting overgrowth themselves o Larger and thicker colonies. octocoral. somewhat slower growing. red crustose alga and a mat of amphipod tubes. by Kenneth Sabens determined. sponge. were usually at the top of the competitive hierarchy ‡ Rapidly growing species were first to colonize bare surface leads to dominance in patches ‡ Food limitations o Sessile species are suspension feeders  Beth Okamura determined food depletion with lateral movement of water across bryozoan colony Bryozoan colony: made up of hundreds of tiny individuals = good example of modular organism .

Subtidal Rocky Reefs Predation and Grazing ‡ Mediates competitive interactions o Create empty space allowing shifts in dominance ‡ Allmon and Sebens studied the effects of an invasive carnivore (Nudibranch Tritonia plebeia) on the rocky reefs of Cape Ann.nearly eliminated them allowing urchins to move onto vertical walls and remove the remaining colonies causing a shift in dominance Octocoral: eight branches on each polyp! Distribution of Tritonia plebeia European Nudibranch: Tritonia plebeia . Massachusetts o Nudibranchs feed on Octocoral.

Tritonia plebeia: Nudibranch swimming Nudibranch eating a tube anemone .Subtidal Rocky Reefs European Nudibranch.

Kelp Forests Kelp forest (textbook definition): A shallow. subtidal community in relatively cold water that is dominated by kelps .

Australia.Kelp Forests ‡ Found in shallow. both coasts of South America. New Zealand. open coastal waters o Dependence upon light for photosynthesis restricts kelp to clear. and South Africa  Laminaria: Alaska and northwestern Atlantic More diversity in Pacific than Atlantic! Distribution of Laminariales . America. shallow water o Rarely found deeper than 5-15 m ‡ Relatively cold o Larger kelp forests are restricted to temperatures under 20 degrees Celsius ‡ Found worldwide  Extending to both the Arctic and Antarctic circles  Macrocystis: west coasts of N.

g. Pacific Coast from California to Alaska .Kelp Forests Laminaria ‡ Dominated by brown seaweeds of the group Laminariales o Includes Macrocystis and Laminar ia  Laminaria: relatively simple. Atlantic beds of Nova Scotia  Macrocystis: vast stands of plants that extend from holdfasts on the deep bottom to blades floating at the surface at ~25 m  e. single-bladed seaweeds of a Macrocystis few meters in depth  e.g.

Kelp Forests ‡ Incredible growth rates o Macrocystis: can grow 30 cm per day and 25 m in 120 days .

the life cycle of Laminaria! o Gametophyte . providing greater access to sunlight for Figure 15.Kelp Forests Complex life cycle ‡ Alternates between large asexual sporophyte and tiny filamentous gametophyte o Sporophyte: the "kelp"  Holdfast: part that attaches to the bottom  Stipe: strong and flexible and behaves like a stem  Blade: leaflike portion that is the site of photosynthesis and growth  Tips of blades erode rapidly but growth as the base replaces these tissues  Pneumatocyst (air bladder): gasfilled sacs in algae that cause the algae·s blades to float up toward the water·s surface.9 in your textbook also depicts photosynthesis.

Kelp Forests Macrocystis life cycle: .

potentially shading lower layers near the seafloor .Kelp Forests ‡ Laminaria: stipe is the only support for the plant ‡ Macrocystis: stipe if very long which allows pneumatocysts (large floating air bulbs) to form a canopy above the seabed o Canopy (textbook definition): a layer of photosynthetic organisms high off the water column.

lobsters. etc.Kelp Forests Biological Diversity within Kelp Forests ‡ Kelp forests have several distinct canopy layers (comparable to terrestrial habitats) o Each canopy dominated by a different kelp species ‡ Grazing species common in rock crevices and open areas o Luminaria kelp beds: urchins. limpets. mussels. o Macrocystis kelp beds: abalones. etc. sea urchins. .

Kelp Forests Urchins ‡ Trap waterborne kelp and other seaweed fragments in dorsal spines and tube feet ‡ Transport this material to oral surface and then ingest ‡ Relatively inactive and wait for food to come to them .

Kelp Forests Dense Kelp Forests ‡ Kelp erodes and separates into small particles o This supports populations of suspension feeders (a. filter feeders: feed on small particles that are suspended in the water)  e.g.a.k. mussels ‡ Areas with steep slopes o Particles are transported to deeper waters  Provides food to urchins .

Kelp Forests ‡ Stellar's Sea Cow o Inhabited North Pacific Rim o Browser on the upper kelp canopy o Extinct due to over-hunting by humans (18th century) .

and fish o Dives to bottom. removes an animal. and brings it to the surface to eat  Smashes prey open with a rock and then floats on its back on the surface while eating o Nearly driven to extinction due to hunting for fur  Primarily in northern Gulf of Alaska o Protected in 1911  Rapid population increases both naturally and through reintroductions  Conservation success story Importance: sea otter is a keystone species.g.Kelp Forests ‡ Sea Otter o Major predator on sea urchins. sea otters keep urchin populations "in check") . mollusks. a major top-down driving force of trophic cascade in eastern Pacific kelp communities o Trophic cascade: when predators suppress abundance of prey (e.

.Kelp Forests Sea Otters (Continued..) .

Factors Affecting Kelp Communities ‡ Three Main Factors Affecting Kelp Communities o o o Storms Behavioral changes in herbivores Presence or absence of sea otters 3 1 2 .

Factors Affecting Kelp Communities 1‡ Storms Storms coming from the open sea can rip kelp from their holdfasts o Lower-canopy kelps survive better o Direct impact of storms on kelp Effects of storms over time .

the preferred food after drift algae are no longer available  Resulting in coralline algal turfs and barrens  Coralline algal turfs (textbook definition): a bottom dominated by layers of one or more species of Red algae that secrete a skeleton of calcium carbonate  Barrens (textbook definition): bare bottoms Coralline algae .Factors Affecting Kelp Communities 2‡ Behavioral changes in herbivores o Change urchin behavior to feed on newly recruiting seaweed.

Purple urchins overgraze kelp 3. Barrens are all that is left . Coralline algal turf created 1. Healthy kelp forest This is an unhealthy kelp forest ecosystem! 4.Factors Affecting Kelp Communities 2.

est.Factors Affecting Kelp Communities Case Study ‡ Took place from 1982-2000 ‡ Two study sites: o Marine reserve at Anacapa Island in southern California. fishing greatly reduced population numbers of spiny lobsters (greatest sea urchin predator) o Caused urchins to be 13X more abundant than in protected areas  Caused reductions in kelp because urchins feed primarily on kelp ‡ In protected areas. unprotected fished areas just outside the reserve Southern California Spiny Lobster Panulirus interruptus ‡ In unprotected areas. by the National Park Service in 1978 and protected ever since o Nearby. no fishing caused lobster populations to become 6X more abundant o Lobsters were able to keep urchin populations "in-check" o Kelp forests remained productive and flourished .

kelp forests in the reserve may be more resilient to climatic stress than kelp forests in unprotected sites. some unprotected sites became barren o Protected sites remained healthy ‡ Throughout study: o Unprotected area alternated between kelp forest and barrens o Protected area remained healthy kelp forest Discussion ‡ ".." ..Factors Affecting Kelp Communities Results ‡ 1970s. both sites had healthy kelp forests ‡ After 1982-1983 El Nino.

principal grazers in the system  Sea otter predation lowered sea urchin population densities by 50 percent in the Aleutian Islands .Factors Affecting Kelp Communities 3‡ Presence or absence of sea otters o o "Sea otters sit atop a strongly linked trophic cascade" Prey upon urchins and abalones.

788 otters were dying each year in Kuluk Bay ‡ Shift to otters probably caused by a decline in harbor seals and Steller's sea lions o Caused by a decline in North Pacific fish stocks due to strong fishery pressure Surprise ending! .Factors Affecting Kelp Communities New Trophic Level: ‡ 1990's attacks by killer whales on sea otters were observed regularly in Alaskan coastal waters o In Kuluk Bay otter populations declined 76 percent due to killer whale predation  A killer whale would have to consume 1.825 sea otters each year to maintain its metabolic requirements  It was estimated that 6.

.): ‡ Decline in otter populations have increased sea urchin density o Kelp grazing increased over 90% ..Factors Affecting Kelp Communities New Trophic Level (Continued.

Factors Affecting Kelp Communities Diversity of Carnivores: ‡ Shown to have a positive correlation with kelp biomass and a negative correlation with herbivore abundance o A starfish (Pycnopodia helianthoides) and several species of crabs prey upon purple sea urchins off the coast of California ‡ Carnivore diversity causes top-down indirect effect on kelp growth .

growth. and reproduction  Macrocystis pyrifera and Nereocystis luetkeana grow the large stipe off from the bottom its more vulnerable fronds are floating at the surface and immune from urchin grazing .Factors Affecting Kelp Communities Kelp Resistance to Herbivory: ‡ Kelp closer to the shore (Laminaria) are not mechanically resistant to grazing o Utilize predators of grazers o Rapid colonization.

Factors Affecting Kelp Communities Kelp Resistance to Herbivory: ‡ Deeper water kelp developed chemical mechanisms o Agarum fimbriatum has high concentrations of polyphenolics (chemical) to make the fronds taste bad to grazers o Desmarestia synthesizes sulfuric acid that can erode the Aristotle's lantern of sea urchins (chewing organ) ‡ Lack of grazer predators Aristotle's Lantern Desmarestia Agarum fimbriatum .

. ‡ Rocky reefs are grazed more intensely. . ‡ Rocky reefs often are very patchy. on horizontal benches. with alternations of rocks dominated by rich invertebrate assemblages and turf-forming calcareous red algae. ‡ Subtidal rock wall patches of animals often are short on space. ‡ Rocky subtidal reefs harbor abundant communities of algae and invertebrates and are often dominated by colonial invertebrates. suggesting the importance of competition.. mainly by sea urchins. ‡ Kelp forest-rocky reefs are often dominated in shallow waters by kelps and seaweeds and by epifaunal animals in deeper waters.In Summary (from textbook).

disturbance. a larger diversity of predators beyond sea otters exerts top-down effects. presence or absence of sea otters. ‡ Kelp forests are biologically diverse and support many plant and animal species. ‡ Predation from an offshore source has introduced a new trophic level to some eastern Pacific kelp forests.. and behavioral changes in herbivores. ‡ Kelp communities are often strongly affected by a combination of storms. ‡ Kelps vary widely in their susceptibility to grazing.In Summary (from textbook).. ‡ Kelp forests are dominated by a few species of brown seaweeds of the group Laminariales with fantastic growth rates. . ‡ In lower-latitude California kelp forests. and competition for light. ‡ In kelp forests. succession depends upon the interplay of grazing pressure.

Thanks for visiting Bikini Bottom! One of the best kelp forests around! Any questions or comments?? .

University of Maryland: Center for Environmental Science. p 511-520. Kelp Forests." Marine Education Society of Australia. "Rocky Reefs. 3rd ed. Ventura.Works Cited Behrens MD. Adrian.edu>. <http://www. .edu. MESA. Rocky Reefs. Sixth California Islands Symposium. <http://ian. 2011. "Chapter 15: Sea Grass Beds." Marine Biology. and Coral Reefs.umces. "Temperate Coastal Ecosystem and Habitat Diagrams. Jeffery S. Schwemm CA (eds) Levinton. 6 Mar 2011.au>. 6 Mar 2011. CA." Integration and Application Network. 2011. Lafferty KD. Lafferty KD (2004) Marine Ecology Progress Series 279:129-139 J. Web. Institute for Wildlife Studies. Behrens MD (2005) In: Garcelon DK. (2009): 418-432.mesa. Web.