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13 Benthic Life Habits Notes for Marine Biology: Function, Biodiversity, Ecology By Jeffrey S. Levinton Important Benthic Lifestyles • • • • • • • Benthos Epibenthic Burrowers Borers Infaunal, semi-infaunal Benthic swimmers Interstitial Benthos - Size Classification • Macrobenthos - shortest dimension > 0.5 mm • 0.1 mm < Meiobenthos < 0.5 mm, greater • Microbenthos < 0.1 mm Feeding Classification • • • • • Suspension feeders Deposit feeders Herbivores (macroalgae or microalgae) Carnivores Scavengers Life in Mud and Sand Soft sediments are a mixture of inorganic particles, organic particles, and pore water. Mechanical measures of sediment grain size Particle size - measure of current strength median, silt-clay percent (<62mm) 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 - percent of weight of sediment < 62mm Sedimentary Structures Can result from physical processes: Ripple marks on an intertidal sand flat Sedimentary structures Can result from biogenic processes Sediment surface showing fecal casting (left) and Burrow opening (right) of lugworm Arenicola marina Burrowing in Sediment • Burrowers use hydromechanical and mechanical digging mechanisms to move through the sediment • Watery sediments with high silt clay content have thixotropy - but burrowing also propagates cracks in muddy sediments with lower water content - particles coated with organic polymers stick together1. 1. Dorgan et al. 2005 Nature 433:475 Burrowing in Sediment • 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 Lugworm, Arenicola marina Longitudinal muscle contracting Terminal anchor Eversion of pharynx Circular muscle contracting Penetration ancthor Burrowing in Sediment Inarticulate brachiopod Mechanical displacement burrowing Burrowing in Sediment Biogenic structures - burrowing and processing of sediment affects sedimentary structures 1. Burrowing in mud increases water content of sediment 2. Increases grain size (fecal pellets to aggregated fine particles) 3. Alters vertical and 3-D mechanical, chemical structure Burrowing in Sediment 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 (a) polychaete, (b) harpacticoid copepod, (c) gastrotrich, (d) hydroid, and (e) opisthobranch gastropod Depth Burrowed, pelletized layer Water content Soft-sediment microzone Soft-Sediment Microzone • Strong vertical chemical gradients • Gradients strongly affected by biological activity Soft-Sediment Microzone Soft-Sediment Microzone Aerobic bacteria (use oxygen to break down organic substrates) RPD H2S oxidizing bacteria RPD Fermenting bacteria (break down organic cpds. --> alcohols,fatty acids) Sulfate reducing bacteria (reduce SO4 to H2S) Methanogenic bacteria (break down organic cpds. --> methane) Organic Matter in Sediment • Particulate organic matter derives from sedimenting phytoplankton, particulates from seaweeds, sea grasses • Decomposition involves leaching of cellular constituents, fragmentation from mechanical processes and feeding, and decay by microbes 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 (a) the surface tentacle feeder Hobsonia, (b) the within-sediment tentacle-feeding bivalve Yoldia limatula, (c) the surface depositfeeding siphonate bivalve Macoma, (d) the within-sediment-feeding Atlantic polychaete Pectinaria gouldii, (e) the surface-feeding Corophium volutator, and (f) the deep-feeding Arenicola marina feces Burrow opening Surface trace of burrow of the Lugworm, Arenicola marina (Anglesey, North Wales) Arenicola marina burrow cross section Deposit Feeders • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) Deposit Feeders Deposit Feeders • 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 • Microbial stripping hypothesis: deposit feeders are most efficient at digesting and assimilating benthic microbes (diatoms, bacteria) • At some times of year, fresh phytoplankton sinks and is added to the bottom: another source of nonliving food for deposit feeders Deposit Feeders • 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 • 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 • Some deposit feeder guts - surfactant activity emulsify particles Deposit Feeders - Optimal gut passage time Food digested Richer food Gut passage time See Taghon, G. L. 1981, American Naturalist, 118: 202-214 Deposit Feeders - Main Points • Ingest sediment including nonliving particulate organic matter (POM) and microbes • Microbes digested more efficiently than POM, but bacteria not abundant enough to support most macrobenthic deposit feeders • Some deposit feeders have surfactant digestive activity - loosen POM from sediment particles • POM often dominates total organic matter, so poor digestion can still bring a reward • Surface-feeders (better food) vs. deep head-down feeders (more refactory food) • Spring deposition of POM from water column can bring a nutritious source of food in a pulse • Some environments (mangroves) have a continuous and nutritious source of organic matter. Suspension Feeders • Feed on small particles, low Reynolds number within a chamber (e.g. bivalves), higher Re outside • Passive versus active suspension feeders Suspension Feeders (a) the active suspension-feeding parchment worm Chaetopterus; (b) the suspension-feeding polychaete Serpula, which uses ciliary currents to draw particles to tentacles; (c) cross section of a bivalve mollusk, an active suspension feeder (arrows denote ciliated tracts transporting particles); (d) the acorn barnacle Semibalanus balanoides with cirri protruded like a basket Suspension Feeders Encounters of particles with fibers, direct interception the dominant mode Suspension Feeders Bivalve, x-section Polychaete Serpula Active suspension feeders Barnacle Active Suspension Feeders - Issues • Concentration of particles - saturation and even clogging • Selection of high-quality particles • Current velocity and ability to create current, and keep siphon erect Particle Selection by Bivalves Use surgical telescope to look within mantle cavity Fiber-optic light source Endoscope Video connector Oyster Gill of mussel Mytilus edulis, red shows ventral tract, where particles are gathered and transported in mucus Oyster gill lamella Measuring Selectivity Flow Cytometer: Chlorophyll, Phycoerythrin, Particle Size, Index of refraction Experiment to study particle selectivity of Eastern oyster Crassostrea virginica Oysters fed particles 20 microns in size, either algae or poor-quality cellulose-rich particles Question: Can oysters discriminate between particles Procedure: Endoscope used to see results, but particles samples from two ciliated tracts, one where particles are transported to mouth or another where they are transported to palp and rejected