13 Benthic Life Habits Notes for Marine Biology: Function, Biodiversity, Ecology By Jeffrey S. Levinton ©Jeffrey S. Levinton 2001 Important benthic lifestyles • • • • • • • Benthos Epibenthic Burrowers Borers Infaunal, Semi-infaunal Benthic swimmers Interstitial Benthos - size classification • Macrobenthos - shortest dimension < 0.5 mm • Meiobenthos - smaller than 0.5 mm, greater than 0.1 mm • Microbenthos - < 0.1 mm Feeding Classification • • • • • Suspension feeders Deposit feeders Herbivores (macroalgae or microalgae) Carnivores Scavengers Food classification ambiguities • Herbivores could be classified as suspension feeders who feed on planktonic microalgae • Carnivores could be classified as suspension feeders who feed on zooplankton Life in Mud and Sand Life in Mud and Sand Particle size - important parameter Current strength Sorting - variation of current strength, poor versus well sorted sediment Life in Mud and Sand Particle size - measures Median grain diameter - measured usually by sieving and weighing size fractions Silt-clay fraction - % of weight of sediment < 62m Sedimentary Structures Ripple marks on an intertidal sand flat Burrowing in sediment • Burrowers use hydromechanical and mechanical digging mechanisms to move through the sediment • Watery sediments with high silt clay content have thixotropy - as you move through sediment it takes yet less force to continue to move Burrowing in sediment 2 • Hydromechanical burrowing - combines muscle contraction working against rigid, fluid filled chamber (skeleton) • Form penetration anchor first to allow further extension of body into sediment • Form terminal anchor to allow pulling of rest of body into the sediment Burrowing in sediment -bivalve foot Muscle contraction Longitudinal muscles PA Circular muscles PA = penetration anchor TA = terminal anchor TA Hydromechanical burrowing Lugworm, Arenicola marina Longitudinal muscle contracting Terminal anchor Eversion of pharynx Circular muscle contracting Pentetration ancthor Burrowing in sediment 4 Inarticulate brachiopod burrows by scissoring 2 valves back and forth, which displaces sediment Mechanical displacement burrowing Burrowing in sediment 5 Biogenic structures - burrowing and processing of sediment affects sedimentary structures 1. Burrowing in mud increases water content of sediment 2. Increases grain size (pellets) 3. Alters vertical and 3-D mechanical, chemical structure Burrowing in sediment 8 Biogenic graded bedding Annelid ingests small particles at depth and deposits them on surface Interstitial animals • Live in the pore waters of sediment, usually sand grains • Belong to many taxonomic groups • All share elongate, wormlike form, in order to move through tight spaces Interstitial animals 2 Harpacticoid copepod Polychaete Gastrotrich Hydroid Opisthobranch gastropod Soft-Sediment Microzone • Strong vertical chemical gradients • Gradients strongly affected by biological exchange activity Oxic Redox Potential Discontinuity Anoxic Soft-Sediment Microzone 4 Soft-Sediment Microzone 5 • Microbial types also vary with depth below the sediment water interface • Towards the surface, aerobic bacteria dominate (energetically most efficient to use oxidation) • Deeper, bacteria present that can live in the absence of oxygen and can produce energy through various chemical means Soft-Sediment Microzone 6 Aerobic bacteria (use oxygen to break down organic substrates) RPD RPD Fermenting bacteria (break down organic cpds. --> alcohols,fatty acids) Sulfate reducing bacteria (reduce SO4 to H2S) Methanogenic bacteria (break down organic cpds. --> methane) Benthic Feeding Types Deposit Feeders • Feed upon sediment, within the sediment or at sediment surface • Head-down deposit feeders feed within the sediment at depth, usually on fine particles, defecate at surface • Surface browsers often feed on surface microorganisms such as diatoms Deposit Feeders 2 Surface tentacle feeding polychaete Tentacle feeding bivalve Deep feeding polychaete Surface feeding amphipod Surface siphonate feeding bivalve Deep feeding polychaete feces Burrow opening Surface trace of burrow of the Lugworm, Arenicola marina (Anglesey, North Wales) Arenicola marina burrow cross section Deposit feeders 3 • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) Deposit feeders 4 • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) • In some sediments non-living organic matter is abundant, so even a low assimilation efficiency returns some nutrition for deposit feeders Deposit feeders 5 • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) • At some times of years, fresh phytoplankton sinks and is added to the bottom: another source of non-living food for deposit feeders Deposit feeders 6 • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) • Detritus from seaweeds is probably more digestible than detritus from seagrasses and marsh grasses, which have much relatively indigestible cellulose Deposit feeders 7 • Deposit feeders feed on sedimentary grains, microbial organisms living on particulate organic particles • Feeding activity accelerates microbial attack - grazing stimulates microbial metabolism, results in tearing apart of organic particles Deposit feeders 9 Egestion Transport to surface of deep sediment Microbial consumption POM deposition Free-burrowing bivalve Sediment stirring RPD Stimulation of microbial growth Tube POM, Metal, sulfur, (mechanically, dissolved organic Chemically, carbon exchange Transformed) Metal, sulfur, dissolved organic carbon exchange Ingestion of sediment Suspension Feeders • Feed on small particles, low Reynolds number • Passive versus active suspension feeders Suspension Feeders 2 Bivalve, x-section Polychaete Serpula Barnacle Suspension Feeders 3 Sea squirt Styela montereyensis Passive suspension feeding mechanism Suspension Feeders 4 Motile particle deposition Direct interception Sieving Inertial impaction = fiber Gravitational deposition Encounters of particles with fibers Suspension Feeders 4 Motile particle deposition Direct interception Sieving Inertial impaction = fiber Gravitational deposition Encounters of particles with fibers Carnivores Bivalve Cuspidaria oystercatcher Gastropod Nucella Polychaete Glycera Crab Callinectes sapidus Herbivores Polychaete Nereis vexillosa Parrot fish radula Chiton Urchins Cellulose feeder Bivalve Teredo foot Greatly elongated mantle Obtains nitrogen with symbiotic nitrogen fixing bacteria The End