page_title: Bioturbation
section_title: Functional Groupings
hierarchical_section_title: Bioturbation / Evolution and Ecology / Ecosystem function / Functional Groupings
caption_attribution_description: Deutsch: Wattwürmer (Arenicola marina)
context_page_description: Bioturbation is defined as the reworking of soils and sediments by animals or plants. These include burrowing, ingestion, and defecation of sediment grains. Bioturbating activities have a profound effect on the environment and are thought to be a primary driver of biodiversity. The formal study of bioturbation began in the 1800s by Charles Darwin experimenting in his garden. The disruption of aquatic sediments and terrestrial soils through bioturbating activities provides significant ecosystem services. These include the alteration of nutrients in aquatic sediment and overlying water, shelter to other species in the form of burrows in terrestrial and water ecosystems, and soil production on land.
  Bioturbators are deemed ecosystem engineers because they alter resource availability to other species through the physical changes they make to their environments. This type of ecosystem change affects the evolution of cohabitating species and the environment, which is evident in trace fossils left in marine and terrestrial sediments. Other bioturbation effects include altering the texture of sediments, bioirrigation, and displacement of microorganisms and non-living particles.
context_section_description: Bioturbators have been organized by a variety of functional groupings based on either ecological characteristics or biogeochemical effects. While the prevailing categorization is based on the way bioturbators transport and interact with sediments, the various groupings likely stem from the relevance of a categorization mode to a field of study (such as ecology or sediment biogeochemistry) and an attempt to concisely organize the wide variety of bioturbating organisms in classes that describe their function. Examples of categorizations include those based on feeding and motility, feeding and biological interactions, and mobility modes. The most common set of groupings are based on sediment transport and are as follows: 
  Gallery-diffusers create complex tube networks within the upper sediment layers and transport sediment through feeding, burrow construction, and general movement throughout their galleries. Gallery-diffusers are heavily associated with burrowing polychaetes, such as Nereis diversicolor and Marenzellaria spp.
  Biodiffusers transport sediment particles randomly over short distances as they move through sediments. Animals mostly attributed to this category include bivalves such as clams, and amphipod species, but can also include larger vertebrates, such as bottom-dwelling fish and rays that feed along the sea floor. Biodiffusers can be further divided into two subgroups, which include epifaunal (organisms that live on the surface sediments) biodiffusers and surface biodiffusers. This subgrouping may also include gallery-diffusers, reducing the number of functional groups.
  Upward-conveyors are oriented head-down in sediments, where they feed at depth and transport sediment through their guts to the sediment surface. Major upward-conveyor groups include burrowing polychaetes like the lugworm, Arenicola marina, and thalassinid shrimps.
  Downward-conveyor species are oriented with their heads towards the sediment-water interface and defecation occurs at depth. Their activities transport sediment from the surface to deeper sediment layers as they feed. Notable downward-conveyors include those in the peanut worm family, Sipunculidae.
  Regenerators are categorized by their ability to release sediment to the overlying water column, which is then dispersed as they burrow. After regenerators abandon their burrows, water flow at the sediment surface can push in and collapse the burrow. Examples of regenerator species include fiddler and ghost crabs.
caption: The lugworm, Arenicola marina.
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