Bioturbation

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. 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 (diagenesis), bioirrigation, and displacement of microorganisms and non-living particles. Bioturbation is sometimes confused with the process of bioirrigation, however these processes differ in what they are mixing; bioirrigation refers to the mixing of water and solutes in sediments and is an effect of bioturbation. Walruses, salmon, and pocket gophers are examples of large bioturbators. Although the activities of these large macrofaunal bioturbators are more conspicuous, the dominant bioturbators are small invertebrates, such as earthworms, polychaetes, ghost shrimp, mud shrimp, and midge larvae. The activities of these small invertebrates, which include burrowing and ingestion and defecation of sediment grains, contribute to mixing and the alteration of sediment structure. Bioturbation's importance for soil processes and geomorphology was first realized by Charles Darwin, who devoted his last scientific book to the subject (The Formation of Vegetable Mould through the Action of Worms). Darwin spread chalk dust over a field to observe changes in the depth of the chalk layer over time. Excavations 30 years after the initial deposit of chalk revealed that the chalk was buried 18 centimeters under the sediment, which indicated a burial rate of 6 millimeters per year. Darwin attributed this burial to the activity of earthworms in the sediment and determined that these disruptions were important in soil formation. In 1891, geologist Nathaniel Shaler expanded Darwin's concept to include soil disruption by ants and trees. The term 'bioturbation' was later coined by Rudolf Richter in 1952 to describe structures in sediment caused by living organisms. Since the 1980s, the term 'bioturbation' has been widely used in soil and geomorphology literature to describe the reworking of soil and sediment by plants and animals. The onset of bioturbation had a profound effect on the environment and the evolution of other organisms. Bioturbation is thought to have been an important co-factor of the Cambrian Explosion, during which most major animal phyla appeared in the fossil record over a short time. Predation arose during this time and promoted the development of hard skeletons, for example bristles, spines, and shells, as a form of armored protection. It is hypothesized that bioturbation resulted from this skeleton formation. These new hard parts enabled animals to dig into the sediment to seek shelter from predators, which created an incentive for predators to search for prey in the sediment (see Evolutionary Arms Race). Burrowing species fed on buried organic matter in the sediment which resulted in the evolution of deposit feeding (consumption of organic matter within sediment). Prior to the development of bioturbation, laminated microbial mats were the dominant biological structures of the ocean floor and drove much of the ecosystem functions. As bioturbation increased, burrowing animals disturbed the microbial mat system and created a mixed sediment layer with greater biological and chemical diversity. This greater biological and chemical diversity is thought to have led to the evolution and diversification of seafloor-dwelling species. An alternate, less widely accepted hypothesis for the origin of bioturbation exists. The trace fossil Nenoxites is thought to be the earliest record of bioturbation, predating the Cambrian Period. The fossil is dated to 555 million years, which places it in the Ediacaran Period. The fossil indicates a 5 centimeter depth of bioturbation in muddy sediments by a burrowing worm. This is consistent with food-seeking behavior, as there tended to be more food resources in the mud than the water column. However, this hypothesis requires more precise geological dating to rule out an early Cambrian origin for this specimen. The evolution of trees during the Devonian Period enhanced soil weathering and increased the spread of soil due to bioturbation by tree roots. Root penetration and uprooting also enhanced soil carbon storage by enabling mineral weathering and the burial of organic matter. 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: The evaluation of the ecological role of bioturbators has largely been species-specific. However, their ability to transport solutes, such as dissolved oxygen, enhance organic matter decomposition and diagenesis, and alter sediment structure has made them important for the survival and colonization by other macrofaunal and microbial communities.