Polychaetes are common in most marine habitats and dominate many infaunal communities. Functional guild classification based on taxonomic identity and morphology has linked community structure to ecological function. The functional guilds now include osmotrophic siboglinids as well as sipunculans, echiurans, and myzostomes, which molecular genetic analyses have placed within Annelida. Advances in understanding of encounter mechanisms explicitly relate motility to feeding mode. New analyses of burrowing mechanics explain the prevalence of bilateral symmetry and blur the boundary between surface and subsurface feeding. The dichotomy between microphagous deposit and suspension feeders and macrophagous carnivores, herbivores, and omnivores is further supported by divergent digestive strategies. Deposit feeding appears to be limited largely to worms longer than 1 cm, with juveniles and small worms in general restricted to ingesting highly digestible organic material and larger, rich food items, blurring the macrophage-microphage dichotomy that applies well to larger worms.

Until now, the analysis of burrowing mechanics has neglected the mechanical properties of impeding, muddy, cohesive sediments, which behave like elastic solids. Here we show that burrowers can progress through such sediments by using a mechanically efficient, previously unsuspected mechanism--crack propagation--in which an alternating 'anchor' system of burrowing serves as a wedge to extend the crack-shaped burrow. The force required to propagate cracks through sediment in this way is relatively small: we find that the force exerted by the annelid worm Nereis virens in making and moving into such a burrow amounts to less than one-tenth of the force it needs to use against rigid aquarium walls.

The polychaete Nereis virens burrows through muddy sediments by exerting dorsoventral forces against the walls of its tongue-depressor-shaped burrow to extend an oblate hemispheroidal crack. Stress is concentrated at the crack tip, which extends when the stress intensity factor (K I ) exceeds the critical stress intensity factor (K Ic ). Relevant forces were measured in gelatin, an analog for elastic muds, by photoelastic stress analysis, and were 0.015±0.001·N (mean ± s.d.; N=5). Measured elastic moduli (E) for gelatin and sediment were used in finite element models to convert the forces in gelatin to those required in muds to maintain the same body shapes observed in gelatin. The force increases directly with increasing sediment stiffness, and is 0.16·N for measured sediment stiffness of E=2.7ϫ10 4 ·Pa. This measurement of forces exerted by burrowers is the first that explicitly considers the mechanical behavior of the sediment. Calculated stress intensity factors fall within the range of critical values for gelatin and exceed those for sediment, showing that crack propagation is a mechanically feasible mechanism of burrowing. The pharynx extends anteriorly as it everts, extending the crack tip only as far as the anterior of the worm, consistent with wedge-driven fracture and drawing obvious parallels between soft-bodied burrowers and more rigid, wedge-shaped burrowers (i.e. clams). Our results raise questions about the reputed high energetic cost of burrowing and emphasize the need for better understanding of sediment mechanics to quantify external energy expenditure during burrowing.Supplementary material available online at

Sea ice typically forms at the ocean's surface, but given a source of supercooled water, an unusual form of ice--anchor ice--can grow on objects in the water column or at the seafloor. For several decades, ecologists have considered anchor ice to be an important agent of disturbance in the shallow-water benthic communities of McMurdo Sound, Antarctica, and potentially elsewhere in polar seas. Divers have documented anchor ice in the McMurdo communities, and its presence coincides with reduced abundance of the sponge Homaxinella balfourensis, which provides habitat for a diverse assemblage of benthic organisms. However, the mechanism of this disturbance has not been explored. Here we show interspecific differences in anchor-ice formation and propagation characteristics for Antarctic benthic organisms. The sponges H. balfourensis and Suberites caminatus show increased incidence of formation and accelerated spread of ice crystals compared to urchins and sea stars. Anchor ice also forms readily on sediments, from which it can grow and adhere to organisms. Our results are consistent with, and provide a potential first step toward, an explanation for disturbance patterns observed in shallow polar benthic communities. Interspecific differences in ice formation raise questions about how surface tissue characteristics such as surface area, rugosity, and mucus coating affect ice formation on invertebrates.

Burrowers and borers are ecosystem engineers that alter their physical environments through bioturbation, bioirrigation and bioerosion. The mechanisms of moving through solid substrata by burrowing or boring depend on the mechanical properties of the medium and the size and morphology of the organism. For burrowing animals, mud differs mechanically from sand; in mud, sediment grains are suspended in an organic matrix that fails by fracture. Macrofauna extend burrows through this elastic mud by fracture. Sand is granular and non-cohesive, enabling grains to more easily move relative to each other, and macrofaunal burrowers use fluidization or plastic rearrangement of grains. In both sand and mud, peristaltic movements apply normal forces and reduce shear. Excavation and localized grain compaction are mechanisms that plastically deform sediments and are effective in both mud and sand, with bulk excavation being used by larger organisms and localized compaction by smaller organisms. Mechanical boring of hard substrata is an extreme form of excavation in which no compaction of burrow walls occurs and grains are abraded with rigid, hard structures. Chemical boring involves secretion to dissolve or soften generally carbonate substrata. Despite substantial differences in the mechanics of the media, similar burrowing behaviors are effective in mud and sand.

SUMMARYBurrowing marine infauna are morphologically diverse and range in size over several orders of magnitude. Whilst effects of ontogenetic and morphological differences on running, flying and swimming are relatively well understood, similar analyses of burrowing mechanics and kinematics are lacking. The polychaete Nereis virens Sars extends its burrow by fracture, using an eversible pharynx to exert force on the walls of the burrow. The resulting stress is amplified at the anterior tip of the burrow, which extends when the stress exceeds the fracture toughness of the material. Here we show that the polychaete Cirriformia moorei extends its burrow by a similar mechanism, but by using its hydrostatic skeleton rather than an eversible pharynx. Based on the dimensionless wedge number, which relates work of fracture to work to maintain body shape against the elasticity of sediment, we predicted that smaller worms would exhibit behaviors characteristic of tougher sediments and that scaling of kinematics would reflect decreasing difficulty in fracturing sediment with increasing body size. We found that smaller worms were relatively blunter and thicker, and had a greater variation of thickness than larger worms as they burrowed. Although these kinematic differences increase the stress amplification at the crack tip, smaller worms still generate lower stress intensity factors. The greater relative body thickness and shape changes of smaller worms are consistent with ontogenetic changes in forces exerted by earthworms, and are likely driven by the challenge of exerting enough stress to extend a crack with a small body size. Supplementary material available online at

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Recent work has shown that muddy sediments are elastic solids through which animals extend burrows by fracture, whereas non-cohesive granular sands fluidize around some burrowers. These different mechanical responses are reflected in the morphologies and behaviours of their respective inhabitants. However, Armandia brevis, a mud-burrowing opheliid polychaete, lacks an expansible anterior consistent with fracturing mud, and instead uses undulatory movements similar to those of sandfish lizards that fluidize desert sands. Here, we show that A. brevis neither fractures nor fluidizes sediments, but instead uses a third mechanism, plastically rearranging sediment grains to create a burrow. The curvature of the undulating body fits meander geometry used to describe rivers, and changes in curvature driven by muscle contraction are similar for swimming and burrowing worms, indicating that the same gait is used in both sediments and water. Large calculated friction forces for undulatory burrowers suggest that sediment mechanics affect undulatory and peristaltic burrowers differently; undulatory burrowing may be more effective for small worms that live in sediments not compacted or cohesive enough to extend burrows by fracture.