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BENTHOS
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BENTHOS: DEFINITIONS Epifauna: live on or are associated with the benthic surface Infauna: live within the substrate Microfauna: animals <0.1 mm in size (e.g. protozoa/bacteria) Meiofauna: animals <0.5 mm in size: “interstitial” (e.g. nematodes, sm. amphipods) Macrofauna: animals > 0.5 mm in size: most familiar kinds of animals (crabs, shrimp, starfish and most mollusks)
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Abiotic Factors Affecting Benthos (to 200 m depth))
Wave action: influence distribution of sediments and physically affect animals Sediments: vary according to wave action (particle size sorting): terrigenous and marine origin (“allochthonous” and “autochthonous”): fine clays go to deeps Salinity and temperature: FW influences; more thermal variability
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Distribution and biomass of benthos
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The Intertidal: Where the Benthos is Most Abundant
Biomass in intertidal= 10X that of 200 m depth and several thousand that of abyss! Not without a cost: wave shock; desiccation; cold; osmotic issues; and land predators. But at high tide: plenty of O2; nutrients; light; and wastes washed away. More relief and habitat diversity= more species diversity
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Reproduction and Dispersal
Broadcast spawning vs. brooding- varying amounts of energy invested, and value of dispersal Where to settle? 1) chemical attractants: settle near your own kind 2) bottom types: settle in appropriate substrates
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PATTERNS OF DIVERSITY WITH DEPTH
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Where the food comes from
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Effects of predator exclusion on the abundance of macrofaunal molluscs, worm and crustaceans
General results: cages have up to 500 x density more infaunal spp. in cages no dominance by any single species
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Soft Sediment Communities
Types of soft-bottom habitats Role of disturbance in regulating community structure Effects of predation, competition and facilitation
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Submarine canyons Latitudinal Diffs. Temp. = sand Tropic. = mud Polar. = Gravel (Arctic w/ riverine mud) Shallow water/Shelf Deep seafloor
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Sandy shores/beaches
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Muddy shores/bays, estuaries, and lagoons
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Nearshore benthic habitats (0-200 m)
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Meiofauna (few mm) Benthic diatoms Harpacticoid copepods
Foraminiferans
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Macrofauna (mm-cm) polychaete worms crustaceans
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Macrofauna (mm-cm) heart urchins pycnogonids brittle stars bivalves
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High biodiversity that varies with depth, sediment type and biotic factors
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Infaunal community “Patchiness” is the rule
1. Biotic interactions: predation, competition, & facilitation 2. Physical factors: disturbance (biotic, physical, and anthropogenic)
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Community patterns and structure
Temperate/tropical Polar
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Megafauna (cm-m) grey whales
walrus Predators have a big effect on community composition
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Caging Studies
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Direct and indirect effects of predation in soft-sediment food webs
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Important classification for understanding effects of disturbance
Life-history groups Capitella captitata Succession
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BIOTURBATION Upogebia- another burrowing shrimp Upogebia BURROW
fecal strands from polychaetes Burrows of Callianassa BURROWING SHRIMP Callianassa
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MORE BIOTURBATORS Harpacticoid copepod Burrowing holothurian
Polychaete: Nereis Oligochaete: Paranais
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The lugworm (Arenicola) and its burrow/fecal castings
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Gastropod: Hydrobia Gastropod 2: Ilyanassa 29
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Sediment modifiers Ammensalism/mutualism Facilitation
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Competition can be important in soft-sediment communities
Competition in a 3-d environment: rarely for space Competition usually for food with big effects on growth, reproduction, and survival. Density-dependence common Competition has a big effect on community structure- depth distribution, population distribution, abundance, and dynamics
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The intermediate disturbance
hypothesis
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Would you expect the intermediate disturbance hypothesis to explain diversity patterns in soft sediments?
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Types and scales of disturbances
in soft-sediments
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Disturbance caused by eutrophication
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Iceberg scour disturbance
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On frequently scoured seafloor, what functional groups would you expect?
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Re-colonization Different mechanisms: Vegetative regrowth of survivors
Recruitment from propagules (including spore and seed bank) Influence of patch characteristics: Size and shape Substrate characteristics (e.g. rock or sediment types, topographic complexity, biogenic structures) Patch location (environmental conditions and proximity to propagule sources) Timing of patch creation (availability of propagules and differences in conditions)
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PHYSICAL DISTURBANCES
Agent of disturbance Waves and currents Water-borne material (sediment, logs, rocks) Ice Direct impacts on organisms and Substrate Sessile organisms detached or broken Mobile animals displaced, injured, or killed Substrate overturned Sediment eroded Organisms abraded, buried, crushed or detached Organisms abraded, detached Sediment and organisms excavated and displaced Habitat or assemblages effected Most, declines with depth Most Rocky intertidal and subtidal, Soft sediment, Seagrass beds, Salt mashes (high lat)
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PHYSICAL DISTURBANCES
Agent of disturbance Extended aerial exposure Temperature extremes Salinity stress and freshwater flooding Anoxia Direct impacts on organisms and Substrate Organisms injured or killed by desiccation, heat, UV by heat or cold. Bleaching by osmotic stress by metabolic stress Habitat or assemblages effected Rocky intertidal Coral reefs Seagrass beds Tide pools, Kelp forests, Coral reefs Rocky intertidal, Salt marsh, Coral reef, Mangrove, Soft sediment Soft sediment, estuaries, semienclosed seas
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PHYSICAL DISTURBANCES
Agent of disturbance Landslides, tectonic events Lava flow, volcanic ash Fire, lightening strikes, Meteorite impacts Direct impacts on organisms and Substrate Organisms abraded, crushed, displaced, or smothered Organisms injured or killed by lava, smothered by ash by heat Direct impact and climate change Habitat or assemblages effected Rocky intertidal and subtidal, Soft sediment, slope and rise,vents Rocky intertidal and subtidal, Seagrass beds, Coral reefs, vents Salt marsh, Mangrove Global (mass extinctions)
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BIOLOGICAL DISTURBANCES
Agent of disturbance Accumulation of plant or animal material (wrack and carcasses) Algal whiplash Bioturbation Sediment excavation by predators Direct impacts on organisms and Substrate Organisms smothered, buried and shaded, chemistry Organisms abraded, recruits vulnerable Organisms buried, sediment load interferes with feeding Organisms displaced, uprooted, and buried Accumulation of debris Habitat or assemblages effected Salt marsh, Seagrass beds, Soft sediment Rocky intertidal and subtidal Soft sediment, Seagrass beds Soft sediments Seagrass beds
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BIOLOGICAL DISTURBANCES
Agent of disturbance Haul out, trampling Red tide Direct impacts on organisms and Substrate Organisms smothered, buried, smashed Organisms suffocated and poisoned Habitat or assemblages effected Rocky intertidal Soft sediment, coastal environments
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Anthropogenic disturbances in soft sediment habitats
Oil spills
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14 years of intensive study
Exxon-Valdez oil spill Bleigh Reef in NPWS 42 million L of oil 1990 km of coastline and 750 km southward 14 years of intensive study Ecosystem level impacts Prince William Sound, AK on 24 March 1989
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Ecological impacts of the EVOS
New understanding of long-term effects and recovery processes. Ecosystem-based toxicology Before: short-term impact assessments and lab studies of toxic effects First real ecosystem impact study General conclusions: 1) Oil persisted beyond a decade in surprising amounts and toxic forms because of the presence of soft-sediments 2) Oil significantly bioavailable to induce chronic biological exposure 3) Longterm effects at population level
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Ecological impacts of the EVOS
3 major pathways of exposure & induction: Acute exposure to oil during spill and subsequent negative health effects Chronic persistence of oil, bio-exposure, and population impacts to species closely associated with shallow sediments Indirect effects related to predator-prey relationships, loss of habitat
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Acute effects of spill Oil in fur, feathers, and ingested
sea otters ~250,000 seabirds 302 harbor seals Oil in fur, feathers, and ingested Mass mortality of macroalgae, benthic invertebrates on shore from a combination toxicity, smothering, and physical displacement caused by high pressure clean-up
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Persistence of oil 40-45% oil grounded in 1989 on 787 km of PWS beaches 7-11% contaminated 1203 km of Gulf of AK coastline 2% remained on beaches after 3.5 yrs (-0.87 per yr) Rates of dispersion and degradation diminished through time Suppressed by physical barriers to disturbance, oxygenation and photolysis Oil trapped in sediments and mussel beds
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Effects of chronic exposure
Chronic exposure of sediment affiliated species Fish embryos exposed to partially weathered oil Multi-ringed PAHs toxic to pink salmon eggs at 1 ppb when exposed for months Toxic to herring eggs when exposed for 16 days Reduced salmon and herring reproduction in many areas
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Led to death from compromised health, growth, or reproduction
Effects of chronic exposure Lower growth rates of salmon = reduced survival Abnormal growth in herring and salmon caused by endocrine disruptors = less fat Led to death from compromised health, growth, or reproduction
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Cascades of indirect effects
Trophic cascades in which predators reduce abundance of their prey which releases the preys food species from control Provision of biogenic habitat by organisms that serve as or creates important physical structure in environment Two most important types of cascades Current Risk Assessment models used for projecting biological injury to marine communities ignore indirect effects
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Four groups of dominant macrofauna in soft bottoms
Class Polychaeta: most numerous: tube-building and burrowing Subphylum Crustacea: ostracods, amphipods, isopods, tanaids, mysids, small decapods Phylum Mollusca: burrowing bivalves and scaphopods, gastropods at surface Phylum Echinodermata: brittle stars, heart urchins, sand dollars, sea cukes
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Soft- and hard-bottom benthic communities
Soft: little ‘relief’: ripple marks, worm tubes, fecal mounds: some differences in sediment grain size: fewer inds. And infauna and more epifauna in sand: more individuals in mud and most are deposit feeders Hard: more ‘relief” and more habitat diversity: increase in suspension feeders
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Feeding strategies Deposit feeders: feed on organically enriched sediments: continuous “reworking” of sediments to extract nutrients: analogous to earthworms: can live in very fine sediments Suspension feeders: filtering devices or mucus nets collect detritus or plankton: need coarser sediments or hard bottom Grazers/predators/scavengers
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Deposit Feeders
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Suspension feeders Bryozoan Hydroid Sponge Bivalve Polychaete Tunicate
Barnacle Amphipod Anemone Brittlestar
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PARALLEL BOTTOM COMMUNITIES
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FUNCTIONAL GROUPS (or: bioturbators)
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FEEDING IN AN INFAUNAL CUCUMBER
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PREDATORS OF BENTHOS
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CAGE STUDIES
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COMPETITION FOR SPACE
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LIFE IN THE MUDS: A COMPLEX SITUATION
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COMMUNITY CHANGES: STORM INDUCED
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SEASONAL POPULATION CHANGE
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LONGER-TERM OSCILLATIONS IN ANOTHER AMPHIPOD
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Biomass of benthos in relation to distance from coast and depth
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RECOVERY OF BENTHIC COMMUNITY
FOLLOWING DEFAUNATION BY RED TIDE
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