20:
475:. Pocket gophers are thought to play an important role in the downslope transport of soil, as the soil that forms their mounds is more susceptible to erosion and subsequent transport. Similar to tree root effects, the construction of burrows-even when backfilled- decreases soil density. The formation of surface mounds also buries surface vegetation, creating nutrient hotspots when the vegetation decomposes, increasing soil organic matter. Due to the high metabolic demands of their burrow-excavating subterranean lifestyle, pocket gophers must consume large amounts of plant material. Though this has a detrimental effect on individual plants, the net effect of pocket gophers is increased plant growth from their positive effects on soil nutrient content and physical soil properties.
589:(area between surface water and groundwater) of rivers and effects the dispersion and retention of marine derived nutrients (MDN) within the river ecosystem. MDN are delivered to river and stream ecosystems by the fecal matter of spawning salmon and the decaying carcasses of salmon that have completed spawning and died. Numerical modeling suggests that residence time of MDN within a salmon spawning reach is inversely proportional to the amount of redd construction within the river. Measurements of respiration within a salmon-bearing river in Alaska further suggest that salmon bioturbation of the river bed plays a significant role in mobilizing MDN and limiting primary productivity while salmon spawning is active. The river ecosystem was found to switch from a net
664:(organic waste). These large quantities, in addition to typically small sediment grain size and dense populations, make bioturbators important in estuarine respiration. Bioturbators enhance the transport of oxygen into sediments through irrigation and increase the surface area of oxygenated sediments through burrow construction. Bioturbators also transport organic matter deeper into sediments through general reworking activities and production of fecal matter. This ability to replenish oxygen and other solutes at sediment depth allows for enhanced respiration by both bioturbators as well as the microbial community, thus altering estuarine elemental cycling.
264:
organic matter decomposition and sediment oxygen uptake. In addition to the effects of burrowing activity on microbial communities, studies suggest that bioturbator fecal matter provides a highly nutritious food source for microbes and other macrofauna, thus enhancing benthic microbial activity. This increased microbial activity by bioturbators can contribute to increased nutrient release to the overlying water column. Nutrients released from enhanced microbial decomposition of organic matter, notably limiting nutrients, such as ammonium, can have bottom-up effects on ecosystems and result in increased growth of phytoplankton and bacterioplankton.
635:
897:, sediments are important for preserving a wide variety of fossils. Evidence of bioturbation has been found in deep-sea sediment cores including into long records, although the act extracting the core can disturb the signs of bioturbation, especially at shallower depths. Arthropods, in particular are important to the geologic record of bioturbation of Eolian sediments. Dune records show traces of burrowing animals as far back as the lower Mesozoic (250 Million years ago), although bioturbation in other sediments has been seen as far back as 550 Ma.
532:(detritus worms) are important agents of bioturbation in these ecosystems and have different effects based on their respective feeding habits. Tubificid worms do not form burrows, they are upward conveyors. Chironomids, on the other hand, form burrows in the sediment, acting as bioirrigators and aerating the sediments and are downward conveyors. This activity, combined with chironomid's respiration within their burrows, decrease available oxygen in the sediment and increase the loss of nitrates through enhanced rates of
422:
225:
647:
238:
915:). 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
517:). The major nutrients of interest in this flux are nitrogen and phosphorus, which often limit the levels of primary production in an ecosystem. Bioturbation increases the flux of mineralized (inorganic) forms of these elements, which can be directly used by primary producers. In addition, bioturbation increases the water column concentrations of nitrogen and phosphorus-containing organic matter, which can then be consumed by fauna and mineralized.
117:
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:
505:
582:) and porosity of the stream bed. In select rivers, if salmon congregate in large enough concentrations in a given area of the river, the total sediment transport from redd construction can equal or exceed the sediment transport from flood events. The net effect on sediment movement is the downstream transfer of gravel, sand and finer materials and enhancement of water mixing within the river substrate.
854:
598:
organic carbon, also attributed to sediment mobilization from salmon redd construction. While marine derived nutrients are generally thought to increase productivity in riparian and freshwater ecosystems, several studies have suggested that temporal effects of bioturbation should be considered when characterizing salmon influences on nutrient cycles.
677:
bioturbation, the effects of bioturbators on denitrification rates have been found to be greater than that on rates of nitrification, further promoting the removal of biologically available nitrogen. This increased removal of biologically available nitrogen has been suggested to be linked to increased rates of
437:, with root growth and stump decay also contributing to soil transport and mixing. Death and decay of tree roots first delivers organic matter to the soil and then creates voids, decreasing soil density. Tree uprooting causes considerable soil displacement by producing mounds, mixing the soil, or inverting
813:. 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.
775:
Parameterization of bioturbation, however, can vary, as newer and more complex models can be used to fit tracer profiles. Unlike the standard biodiffusion model, these more complex models, such as expanded versions of the biodiffusion model, random walk, and particle-tracking models, can provide more
325:
of larvae of conspecifics (those of the same species) and those of other species, as the resuspension of sediments and alteration of flow at the sediment-water interface can affect the ability of larvae to burrow and remain in sediments. This effect is largely species-specific, as species differences
919:
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
696:
Bioturbation by walrus feeding is a significant source of sediment and biological community structure and nutrient flux in the Bering Sea. Walruses feed by digging their muzzles into the sediment and extracting clams through powerful suction. By digging through the sediment, walruses rapidly release
676:
coupling contributes to greater removal of biologically available nitrogen in shallow and coastal environments, which can be further enhanced by the excretion of ammonium by bioturbators and other organisms residing in bioturbator burrows. While both nitrification and denitrification are enhanced by
520:
Lake and pond sediments often transition from the aerobic (oxygen containing) character of the overlaying water to the anaerobic (without oxygen) conditions of the lower sediment over sediment depths of only a few millimeters, therefore, even bioturbators of modest size can affect this transition of
345:
For example, bioturbating animals are hypothesized to have affected the cycling of sulfur in the early oceans. According to this hypothesis, bioturbating activities had a large effect on the sulfate concentration in the ocean. Around the
Cambrian-Precambrian boundary (539 million years ago), animals
116:
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
304:
Bioturbators can also inhibit the presence of other benthic organisms by smothering, exposing other organisms to predators, or resource competition. While thalassinidean shrimps can provide shelter for some organisms and cultivate interspecies relationships within burrows, they have also been shown
248:
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
1542:. Aller, Josephine Y., Woodin, Sarah Ann, 1945–, Aller, Robert C., Belle W. Baruch Institute for Marine Biology and Coastal Research. Columbia, S.C.: Published for the Belle W. Baruch Institute for Marine Biology and Coastal Research by the University of South Carolina Press. 2001. pp. 73–86.
885:
within geology. The study of bioturbator ichnofabrics uses the depth of the fossils, the cross-cutting of fossils, and the sharpness (or how well defined) of the fossil to assess the activity that occurred in old sediments. Typically the deeper the fossil, the better preserved and well defined the
597:
system in response to decreased primary production and increased respiration. The decreased primary production in this study was attributed to the loss of benthic primary producers who were dislodged due to bioturbation, while increased respiration was thought to be due to increased respiration of
512:
The sediments of lake and pond ecosystems are rich in organic matter, with higher organic matter and nutrient contents in the sediments than in the overlying water. Nutrient re-regeneration through sediment bioturbation moves nutrients into the water column, thereby enhancing the growth of aquatic
350:
to overlying water causing sulfide to oxidize, which increased the sulfate composition in the ocean. During large extinction events, the sulfate concentration in the ocean was reduced. Although this is difficult to measure directly, seawater sulfur isotope compositions during these times indicates
275:
burrow and create mounds that have a complex system of air ducts and evaporation devices that create a suitable microclimate in an unfavorable physical environment. Many species are attracted to bioturbator burrows because of their protective capabilities. The shared use of burrows has enabled the
263:
occurring around burrows. As bioturbators burrow, they also increase the surface area of sediments across which oxidized and reduced solutes can be exchanged, thereby increasing the overall sediment metabolism. This increase in sediment metabolism and microbial activity further results in enhanced
359:
by increasing the sequestration of phosphorus above normal chemical rates. The sequestration of phosphorus limits oxygen concentrations by decreasing production on a geologic time scale. This decrease in production results in an overall decrease in oxygen levels, and it has been proposed that the
495:
for moving material between the sediments and water column, feeding on sediment organic matter and transporting mineralized nutrients into the water column. Both benthivorous and anadromous fish can affect ecosystems by decreasing primary production through sediment re-suspension, the subsequent
552:
phytoplankton colonies benefit from both increased suspended nutrients and from recruitment of buried phytoplankton cells released from the sediments by the fish bioturbation. Macrophyte growth has also been shown to be inhibited by displacement from the bottom sediments due to fish burrowing.
551:
by the re-suspension of benthic sediments. This increased turbidity limits light penetration and coupled with increased nutrient flux from the sediment into the water column, inhibits the growth of macrophytes (aquatic plants) favoring the growth of phytoplankton in the surface waters. Surface
539:
The increased oxygen input to sediments by macroinvertebrate bioirrigation coupled with bioturbation at the sediment-water interface complicates the total flux of phosphorus . While bioturbation results in a net flux of phosphorus into the water column, the bio-irrigation of the sediments with
1355:
Michaud, Emma; Desrosiers, Gaston; Mermillod-Blondin, Florian; Sundby, Bjorn; Stora, Georges (2006-10-03). "The functional group approach to bioturbation: II. The effects of the Macoma balthica community on fluxes of nutrients and dissolved organic carbon across the sediment–water interface".
717:
in solute concentration and mineral distribution in the sediment. It has been suggested that higher benthic diversity in the deep sea could lead to more bioturbation which, in turn, would increase the transport of organic matter and nutrients to benthic sediments. Through the consumption of
78:, 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.
280:
between bioturbators and the many species that utilize their burrows. For example, gobies, scale-worms, and crabs live in the burrows made by innkeeper worms. Social interactions provide evidence of co-evolution between hosts and their burrow symbionts. This is exemplified by shrimp-goby
470:
also play an important role in the production of soil, possibly with an equal magnitude to abiotic processes. Pocket gophers form above-ground mounds, which moves soil from the lower soil horizons to the surface, exposing minimally weathered rock to surface erosion processes, speeding
360:
rise of bioturbation corresponds to a decrease in oxygen levels of that time. The negative feedback of animals sequestering phosphorus in the sediments and subsequently reducing oxygen concentrations in the environment limits the intensity of bioturbation in this early environment.
667:
The effects of bioturbation on the nitrogen cycle are well-documented. Coupled denitrification and nitrification are enhanced due to increased oxygen and nitrate delivery to deep sediments and increased surface area across which oxygen and nitrate can be exchanged. The enhanced
206:
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
697:
large amounts of organic material and nutrients, especially ammonium, from the sediment to the water column. Additionally, walrus feeding behavior mixes and oxygenates the sediment and creates pits in the sediment which serve as new habitat structures for invertebrate larvae.
869:, or the study of "trace fossils", which, in the case of bioturbators, are fossils left behind by digging or burrowing animals. This can be compared to the footprint left behind by these animals. In some cases bioturbation is so pervasive that it completely obliterates
829:
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
2942:
Mermillod-Blondin, Florian; Gaudet, Jean-Paul; Gerino, Magali; Desrosiers, Gaston; Jose, Jacques; Châtelliers, Michel Creuzé des (2004-07-01). "Relative influence of bioturbation and predation on organic matter processing in river sediments: a microcosm experiment".
563:
show similar responses to bioturbation activities, with chironomid larvae and tubificid worm macroinvertebrates remaining as important benthic agents of bioturbation. These environments can also be subject to strong season bioturbation effects from anadromous fish.
800:
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
767:
from nuclear fallout, introduced particles including glass beads tagged with radioisotopes or inert fluorescent particles, and chlorophyll a. Biodiffusion models are then fit to vertical distributions (profiles) of tracers in sediments to provide values for
3631:
Gerino, M.; Aller, R.C.; Lee, C.; Cochran, J.K.; Aller, J.Y.; Green, M.A.; Hirschberg, D. (1998). "Comparison of
Different Tracers and Methods Used to Quantify Bioturbation During a Spring Bloom: 234-Thorium, Luminophores and Chlorophylla".
2600:
Granberg, Maria E.; Gunnarsson, Jonas S.; Hedman, Jenny E.; Rosenberg, Rutger; Jonsson, Per (2008). "Bioturbation-driven release of organic contaminants from Baltic Sea sediments mediated by the invading polychaete
Marenzelleria neglecta".
65:
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
297:, meaning their existence depends on the host bioturbator and its burrow. Although newly hatched blind gobies have fully developed eyes, their eyes become withdrawn and covered by skin as they develop. They show evidence of commensal
376:, bioturbating animals can mix the surface layer and cause the release of sequestered contaminants into the water column. Upward-conveyor species, like polychaete worms, are efficient at moving contaminated particles to the surface.
547:. Of the bioturbating, benthivorous fish species, carp in particular are important ecosystem engineers and their foraging and burrowing activities can alter the water quality characteristics of ponds and lakes. Carp increase water
124:
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
3031:
Buxton, Todd H.; Buffington, John M.; Tonina, Daniele; Fremier, Alexander K.; Yager, Elowyn M. (2015-04-13). "Modeling the influence of salmon spawning on hyporheic exchange of marine-derived nutrients in gravel stream beds".
521:
the chemical characteristics of sediments. By mixing anaerobic sediments into the water column, bioturbators allow aerobic processes to interact with the re-suspended sediments and the newly exposed bottom sediment surfaces.
354:
Bioturbators have also altered phosphorus cycling on geologic scales. Bioturbators mix readily available particulate organic phosphorus (P) deeper into ocean sediment layers which prevents the precipitation of phosphorus
3882:
Gingras, Murray; Hagadorn, James W.; Seilacher, Adolf; Lalonde, Stefan V.; Pecoits, Ernesto; Petrash, Daniel; Konhauser, Kurt O. (2011-05-15). "Possible evolution of mobile animals in association with microbial mats".
805:). 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
758:
term. This representation and subsequent variations account for the different modes of mixing by functional groups and bioirrigation that results from them. The biodiffusion coefficient is usually measured using
2846:
Matsuzaki, Shin-ichiro S.; Usio, Nisikawa; Takamura, Noriko; Washitani, Izumi (2007-01-01). "Effects of common carp on nutrient dynamics and littoral community composition: roles of excretion and bioturbation".
1216:
Humphreys, G. S., and
Mitchell, P. B., 1983, A preliminary assessment of the role of bioturbation and rainwash on sandstone hillslopes in the Sydney Basin, in Australian and New Zealand Geomorphology Group, p.
3838:
Rogov, Vladimir; Marusin, Vasiliy; Bykova, Natalia; Goy, Yuriy; Nagovitsin, Konstantin; Kochnev, Boris; Karlova, Galina; Grazhdankin, Dmitriy (2012-05-01). "The oldest evidence of bioturbation on Earth".
3805:
Delmotte, Sebastien; Gerino, Magali; Thebault, Jean Marc; Meysman, Filip J. R. (2008-03-01). "Modeling effects of patchiness and biological variability on transport rates within bioturbated sediments".
42:
by animals or plants. It includes 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
3770:
Wheatcroft, R. A.; Jumars, P. A.; Smith, C. R.; Nowell, A. R. M. (1990-02-01). "A mechanistic view of the particulate biodiffusion coefficient: Step lengths, rest periods and transport directions".
2698:
Reible, D.D.; Popov, V.; Valsaraj, K.T.; Thibodeaux, L.J.; Lin, F.; Dikshit, M.; Todaro, M.A.; Fleeger, J.W. (1996). "Contaminant fluxes from sediment due to tubificid oligochaete bioturbation".
3168:
Henriksen, K.; Rasmussen, M. B.; Jensen, A. (1983). "Effect of
Bioturbation on Microbial Nitrogen Transformations in the Sediment and Fluxes of Ammonium and Nitrate to the Overlaying Water".
3133:
Kristensen, Erik; Jensen, Mikael Hjorth; Andersen, Torben Kjær (1985). "The impact of polychaete (Nereis virens Sars) burrows on nitrification and nitrate reduction in estuarine sediments".
1445:
Josefson, Alf B. (1985-12-30). "Distribution of diversity and functional groups of marine benthic infauna in the
Skagerrak (eastern North Sea) - Can larval availability affect diversity?".
154:
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
2554:
Josefsson, Sarah; Leonardsson, Kjell; Gunnarsson, Jonas S.; Wiberg, Karin (2010). "Bioturbation-driven release of buried PCBs and PBDEs from different depths in contaminated sediments".
402:
organic contaminants into the sediment. Burial of uncontaminated particles by bioturbating organisms provides more absorptive surfaces to sequester chemical pollutants in the sediments.
281:
associations. Shrimp burrows provide shelter for gobies and gobies serve as a scout at the mouth of the burrow, signaling the presence of potential danger. In contrast, the blind goby
433:
from the lower soil depths to the surface. Terrestrial bioturbation is important in soil production, burial, organic matter content, and downslope transport. Tree roots are sources of
2234:
Dumbauld, Brett R; Brooks, Kenneth M; Posey, Martin H (2001). "Response of an
Estuarine Benthic Community to Application of the Pesticide Carbaryl and Cultivation of Pacific Oysters (
108:. 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.
394:
can burrow to 35-50 centimeters which is deeper than native animals, thereby releasing previously sequestered contaminants. However, bioturbating animals that live in the sediment (
92:
are examples of large bioturbators. Although the activities of these large macrofaunal bioturbators are more conspicuous, the dominant bioturbators are small invertebrates, such as
158:(organisms that live on the surface sediments) biodiffusers and surface biodiffusers. This subgrouping may also include gallery-diffusers, reducing the number of functional groups.
164:
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
190:
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
784:
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
3547:
Bunzl, K (2002). "Transport of fallout radiocesium in the soil by bioturbation: a random walk model and application to a forest soil with a high abundance of earthworms".
2891:
Holtgrieve, Gordon W.; Schindler, Daniel E. (2011-02-01). "Marine-derived nutrients, bioturbation, and ecosystem metabolism: reconsidering the role of salmon in streams".
1097:
Wilkinson, Marshall T.; Richards, Paul J.; Humphreys, Geoff S. (2009-12-01). "Breaking ground: Pedological, geological, and ecological implications of soil bioturbation".
911:
3504:
Sharma, P.; Gardner, L. R.; Moore, W. S.; Bollinger, M. S. (1987-03-01). "Sedimentation and bioturbation in a salt marsh as revealed by 210Pb, 137Cs, and 7Be studies12".
3318:
Aller, Robert C.; Hall, Per O.J.; Rude, Peter D.; Aller, J.Y. (1998). "Biogeochemical heterogeneity and suboxic diagenesis in hemipelagic sediments of the Panama Basin".
2790:
Adámek, Zdeněk; Maršálek, Blahoslav (2013-02-01). "Bioturbation of sediments by benthic macroinvertebrates and fish and its implication for pond ecosystems: a review".
3113:
Aller, Robert C. (1988). "13. Benthic fauna and biogeochemical processes in marine sediments: the role of burrow structures". In
Blackburn, T.H.; Sørensen, J. (eds.).
1617:
Branch, G.M.; Pringle, A. (1987). "The impact of the sand prawn
Callianassa kraussi Stebbing on sediment turnover and on bacteria, meiofauna, and benthic microflora".
1136:
Shaler, N. S., 1891, The origin and nature of soils, in Powell, J. W., ed., USGS 12th Annual report 1890–1891: Washington, D.C., Government Printing Office, p. 213-45.
722:(POC) into the sediment where it is consumed by sediment dwelling animals and bacteria. Incorporation of POC into the food webs of sediment dwelling animals promotes
3272:
Morys, C; Forster, S; Graf, G (2016-09-28). "Variability of bioturbation in various sediment types and on different spatial scales in the southwestern Baltic Sea".
726:
by removing carbon from the water column and burying it in the sediment. In some deep-sea sediments, intense bioturbation enhances manganese and nitrogen cycling.
3670:
Reed, Daniel C.; Huang, Katherine; Boudreau, Bernard P.; Meysman, Filip J.R. (2006). "Steady-state tracer dynamics in a lattice-automaton model of bioturbation".
574:(gravel depressions or "nests" containing eggs buried under a thin layer of sediment) in rivers and streams and by mobilization of nutrients. The construction of
4105:
Hertweck, G; Liebezeit, G (2007). "Bioturbation structures of polychaetes in modern shallow marine environments and their analogues to Chondrites group traces".
3590:
Solan, Martin; Wigham, Benjamin D.; Hudson, Ian R.; Kennedy, Robert; Coulon, Christopher H.; Norling, Karl; Nilsson, Hans C.; Rosenberg, Rutger (2004-04-28).
3223:
Danovaro, Roberto; Gambi, Cristina; Dell'Anno, Antonio; Corinaldesi, Cinzia; Fraschetti, Simonetta; Vanreusel, Ann; Vincx, Magda; Gooday, Andrew J. (2008).
301:
because it is hypothesized that the lack of light in the burrows where the blind gobies reside is responsible for the evolutionary loss of functional eyes.
58:
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.
2445:
Dale, A.W (2016). "A model for microbial phosphorus cycling in bioturbated marine sediments: Significance for phosphorus burial in the early Paleozoic".
543:
The presence of macroinvertebrates in sediment can initiate bioturbation due to their status as an important food source for benthivorous fish such as
540:
oxygenated water enhances the adsorption of phosphorus onto iron-oxide compounds, thereby reducing the total flux of phosphorus into the water column.
825:
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
317:. This has become a serious issue in the northwestern United States, as ghost and mud shrimp (thalassinidean shrimp) are considered pests to bivalve
3490:
309:, because thalassinidean shrimps can smother bivalves when they resuspend sediment. They have also been shown to exclude or inhibit polychaetes,
3982:"Terrestrial-marine teleconnections in the Devonian: links between the evolution of land plants, weathering processes, and marine anoxic events"
3592:"In situ quantification of bioturbation using time lapse fluorescent sediment profile imaging (f SPI), luminophore tracers and model simulation"
3413:
Smith, Craig R.; Berelson, Will; Demaster, David J.; Dobbs, Fred C.; Hammond, Doug; Hoover, Daniel J.; Pope, Robert H.; Stephens, Mark (1997).
570:
function as bioturbators on both gravel to sand-sized sediment and a nutrient scale, by moving and re-working sediments in the construction of
734:
The role of bioturbators in sediment biogeochemistry makes bioturbation a common parameter in sediment biogeochemical models, which are often
491:(migrating) fish such as salmon. Anadromous fish migrate from the sea into fresh-water rivers and streams to spawn. Macroinvertebrates act as
446:, such as earth worms and small mammals, form passageways for air and water transport which changes the soil properties, such as the vertical
146:
transport sediment particles randomly over short distances as they move through sediments. Animals mostly attributed to this category include
50:
experimenting in his garden. The disruption of aquatic sediments and terrestrial soils through bioturbating activities provides significant
1802:
Caliman, Adriano; Carneiro, Luciana S.; Leal, João J. F.; Farjalla, Vinicius F.; Bozelli, Reinaldo L.; Esteves, Francisco A. (2012-09-12).
342:, primarily through nutrient cycling. Bioturbators played, and continue to play, an important role in nutrient transport across sediments.
1804:"Community Biomass and Bottom up Multivariate Nutrient Complementarity Mediate the Effects of Bioturbator Diversity on Pelagic Production"
841:
enhanced soil weathering and increased the spread of soil due to bioturbation by tree roots. Root penetration and uprooting also enhanced
429:
Plants and animals utilize soil for food and shelter, disturbing the upper soil layers and transporting chemically weathered rock called
3191:
Kristensen, Erik (1985). "Oxygen and Inorganic Nitrogen Exchange in a "Nereis virens" (Polychaeta) Bioturbated Sediment-Water System".
1598:
Andersen, FO; Kristensen, E (1991). "Effects of burrowing macrofauna on organic matter decomposition in coastal marine sediments".
969:
Ray, G. Carleton; McCormick-Ray, Jerry; Berg, Peter; Epstein, Howard E. (2006). "Pacific walrus: Benthic bioturbator of Beringia".
2095:
Rhoads, DC; Young, DK (1970). "The influence of deposit-feeding organisms on sediment stability and community trophic structure".
19:
1871:"Sediment-to-water fluxes of particulate material and microbes by resuspension and their contribution to the planktonic food web"
487:(bottom-dwelling) fish, macroinvertebrates such as worms, insect larvae, crustaceans and molluscs, and seasonal influences from
2140:
Pillay, D.; Branch, G. M.; Forbes, A. T. (2007-09-01). "Experimental evidence for the effects of the thalassinidean sandprawn
1296:
Braeckman, U; Provoost, P; Gribsholt, B; Gansbeke, D Van; Middelburg, JJ; Soetaert, K; Vincx, M; Vanaverbeke, J (2010-01-28).
796:
arose during this time and promoted the development of hard skeletons, for example bristles, spines, and shells, as a form of
1391:
Weinberg, James R. (1984-08-28). "Interactions between functional groups in soft-substrata: Do species differences matter?".
1956:
Eisenberg, John F.; Kinlaw, Al (1999). "Introduction to the Special Issue: ecological significance of open burrow systems".
893:
from bioturbation have been found in marine sediments from tidal, coastal and deep sea sediments. In addition sand dune, or
413:
is still affected by bioturbation in the modern Earth. Some examples in the terrestrial and aquatic ecosystems are below.
1654:"Biodiversity of benthic microbial communities in bioturbated coastal sediments is controlled by geochemical microniches"
2118:
3466:
1999:
1547:
1538:"A new model of bioturbation for a functional approach to sediment reworking resulting from macrobenthic communities".
3075:
2284:
Olivier, Frédéric; Desroy, Nicolas; Retière, Christian (1996). "Habitat selection and adult-recruit interactions in
4162:
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496:
displacement of benthic primary producers, and recycling nutrients from the sediment back into the water column.
2498:
70:
left in marine and terrestrial sediments. Other bioturbation effects include altering the texture of sediments (
4182:
4147:
739:
660:, such as estuaries, are generally highly productive, which results in the accumulation of large quantities of
1332:
743:
714:
326:
in resuspension and burrowing modes have variable effects on fluid dynamics at the sediment-water interface.
2740:
Reichman, O.J.; Seabloom, Eric W. (2002). "The role of pocket gophers as subterranean ecosystem engineers".
2325:"Influence of adult density on recruitment into soft sediments: a short-term in situ sublittoral experiment"
372:
of contaminants from the sediment to the water column, depending on the mechanism of sediment transport. In
293:
shrimp burrows where there is not much light. The blind goby is an example of a species that is an obligate
1761:
Kristensen, Erik; Delefosse, Matthieu; Quintana, Cintia O.; Flindt, Mogens R.; Valdemarsen, Thomas (2014).
1711:"Aerobic Decomposition of Sediment and Detritus as a Function of Particle Surface Area and Organic Content"
735:
454:, and nutrient content. Invertebrates that burrow and consume plant detritus help produce an organic-rich
776:
accuracy, incorporate different modes of sediment transport, and account for more spatial heterogeneity.
3936:
Brasier, Martin D.; McIlroy, Duncan; Liu, Alexander G.; Antcliffe, Jonathan B.; Menon, Latha R. (2013).
3459:
Diagenetic Models and Their Implementation : Modelling Transport and Reactions in Aquatic Sediments
3099:
2023:
1571:
719:
718:
surface-derived organic matter, animals living on the sediment surface facilitate the incorporation of
607:
447:
252:
Microbial communities are greatly influenced by bioturbator activities, as increased transport of more
713:. In low energy regions (areas with relatively still water), bioturbation is the only force creating
682:
3905:
2039:"Macrofauna associated with echiuran burrows: a review with new observations of the innkeeper worm,
4167:
1165:
Kristensen, E; Penha-Lopes, G; Delefosse, M; Valdemarsen, T; Quintana, CO; Banta, GT (2012-02-02).
356:
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to have strong negative effects on other species, especially those of bivalves and surface-grazing
187:
1994:. Lacey, Eileen A., Patton, James L., Cameron, Guy N. Chicago: University of Chicago Press. 2000.
267:
Burrows offer protection from predation and harsh environmental conditions. For example, termites
298:
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Taylor, A. M.; Goldring, R. (1993). "Description and analysis of bioturbation and ichnofabric".
1763:"Influence of benthic macrofauna community shifts on ecosystem functioning in shallow estuaries"
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2119:"The effects of bioturbation on the infaunal community adjacent to an offshore hardbottom reef"
1033:
Meysman, F; Meddelburg, J; Heip, C (2006). "Bioturbation: a fresh look at Darwin's last idea".
870:
802:
579:
380:
animals can remobilize contaminants previously considered to be buried at a safe depth. In the
338:
Since its onset around 539 million years ago, bioturbation has been responsible for changes in
270:
3001:
Gottesfeld, AS; Hassan, MA; Tunnicliffe, JF (2008). "Salmon bioturbation and stream process".
3705:
Aquino, Tomás; Roche, Kevin R.; Aubeneau, Antoine; Packman, Aaron I.; Bolster, Diogo (2017).
651:
322:
2499:"Stabilization of the coupled oxygen and phosphorus cycles by the evolution of bioturbation"
1298:"Role of macrofauna functional traits and density in biogeochemical fluxes and bioturbation"
321:
operations. The presence of bioturbators can have both negative and positive effects on the
4114:
4041:
3949:
3892:
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3718:
3679:
3641:
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3556:
3513:
3426:
3369:
3327:
3281:
3236:
3225:"Exponential Decline of Deep-Sea Ecosystem Functioning Linked to Benthic Biodiversity Loss"
3142:
2952:
2900:
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2707:
2662:
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2297:
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1965:
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was first realized by Charles Darwin, who devoted his last scientific book to the subject (
755:
723:
260:
2288:(Malmgren) (Annelida: Polychaeta) post-larval populations: Results of flume experiments".
1244:"Expanding the envelope: linking invertebrate bioturbators with micro-evolutionary change"
8:
2860:
816:
An alternate, less widely accepted hypothesis for the origin of bioturbation exists. The
634:
434:
131:
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3683:
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1969:
1926:
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1819:
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1630:
1497:
1404:
1369:
1313:
1259:
1182:
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1046:
982:
4172:
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Ahlbrandt, T. S.; Andrews, S.; Gwynne, D.T. (1978). "Bioturbation in eolian deposits".
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4014:
3981:
3787:
3747:
3706:
3484:
3395:
3200:
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2419:
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2017:
1938:
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1803:
1565:
1167:"What is bioturbation? The need for a precise definition for fauna in aquatic sciences"
785:
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706:
62:
51:
3568:
3439:
3414:
3339:
2753:
2309:
2259:
2070:
1426:
Fauchald, K; Jumars, P (1979). "Diet of worms: a study of polychaete feeding guilds".
709:
functioning depends on the use and recycling of nutrients and organic inputs from the
681:
in microenvironments within burrows, as indicated by evidence of nitrogen fixation by
27:. Walrus bioturbations in Arctic benthic sediments have large-scale ecosystem effects.
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4001:
3918:
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3791:
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2005:
1995:
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678:
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514:
253:
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The construction of salmon redds increases sediment and nutrient fluxes through the
4122:
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3957:
3910:
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1973:
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230:
3070:. American Geophysical Union. Washington, D.C.: American Geophysical Union. 2005.
4126:
1828:
809:
were the dominant biological structures of the ocean floor and drove much of the
673:
611:
560:
533:
529:
492:
339:
256:
3986:
Philosophical Transactions of the Royal Society of London B: Biological Sciences
1377:
990:
571:
3819:
3783:
3730:
3358:"Climate variation, carbon flux, and bioturbation in the abyssal North Pacific"
2195:
Tamaki, Akio (1988). "Effects of the bioturbating activity of the ghost shrimp
1054:
586:
575:
488:
472:
467:
459:
347:
330:
bioturbators may also hamper recruitment by consuming recently settled larvae.
327:
89:
47:
3691:
3525:
3382:
3357:
3249:
3224:
2811:
2466:
2165:
1735:
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4156:
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3922:
3868:
3738:
3533:
3476:
3391:
3301:
3085:
3053:
2972:
2920:
2819:
2410:
2377:"Animal evolution, bioturbation, and the sulfate concentration of the oceans"
2173:
1837:
1788:
1779:
1762:
1744:
1687:
1466:
1341:
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484:
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177:
137:
101:
75:
3045:
2401:
2009:
1913:
Hansell, M. H. (1993). "The Ecological Impact of Animal Nests and Burrows".
1557:
1482:"Influence of relative mobilities on the composition of benthic communities"
259:, such as oxygen, to typically highly reduced sediments at depth alters the
3997:
3756:
3653:
3576:
3258:
2928:
2684:
2630:
2583:
2428:
2267:
1977:
1855:
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890:
882:
817:
525:
463:
438:
294:
208:
195:
105:
67:
43:
1678:
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398:) can also reduce the flux of contaminants to the water column by burying
3356:
Vardaro, Michael F.; Ruhl, Henry A.; Smith, Kenneth Jr. L. (2009-11-01).
1164:
894:
845:
storage by enabling mineral weathering and the burial of organic matter.
842:
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97:
71:
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2675:
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2622:
2575:
1322:
1297:
1268:
1243:
1191:
1166:
920:
to describe the reworking of soil and sediment by plants and animals.
3914:
3860:
3222:
2525:
2117:
Dahlgren, Craig P.; Posey, Martin H.; Hulbert, Alan W. (1999-01-01).
2071:"The association between gobiid fishes and burrowing alpheid shrimps"
939:
866:
826:
821:
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793:
590:
548:
430:
314:
277:
191:
155:
151:
93:
2553:
1934:
838:
661:
623:
483:
Important sources of bioturbation in freshwater ecosystems include
147:
55:
39:
3068:
Interactions between macro- and microorganisms in marine sediments
2038:
1295:
2599:
1760:
797:
619:
455:
395:
310:
169:
614:, however, they are dominated by small invertebrates, including
3881:
789:
567:
504:
351:
bioturbators influenced the sulfur cycling in the early Earth.
241:
85:
81:
24:
2845:
3804:
3030:
2697:
853:
912:
The Formation of Vegetable Mould through the Action of Worms
3769:
3503:
3000:
1992:
Life underground : the biology of subterranean rodents
1801:
968:
369:
35:
3935:
3704:
3419:
Deep Sea Research Part II: Topical Studies in Oceanography
3412:
1096:
750:, or the biodiffusion coefficient, and is described by a
46:. The formal study of bioturbation began in the 1800s by
4142:
3669:
3589:
3167:
3132:
834:
to rule out an early Cambrian origin for this specimen.
3938:"The oldest evidence of bioturbation on Earth: COMMENT"
3707:"A Process-Based Model for Bioturbation-Induced Mixing"
3320:
Deep Sea Research Part I: Oceanographic Research Papers
4077:
3980:
Algeo, Thomas J.; Scheckler, Stephen E. (1998-01-29).
3837:
3630:
1032:
642:
in the sediment at the bottom of a coastal ecosystems
2283:
2116:
861:
Patterns or traces of bioturbation are preserved in
2890:
2375:Canfield, Donald E.; Farquhar, James (2009-05-19).
2233:
3461:. Berlin, Heidelberg: Springer Berlin Heidelberg.
3317:
3135:Journal of Experimental Marine Biology and Ecology
3034:Canadian Journal of Fisheries and Aquatic Sciences
2651:"The bioturbation-driven chemical release process"
2322:
2201:Journal of Experimental Marine Biology and Ecology
2139:
1652:Bertics, Victoria J; Ziebis, Wiebke (2009-05-21).
1619:Journal of Experimental Marine Biology and Ecology
1597:
1393:Journal of Experimental Marine Biology and Ecology
1358:Journal of Experimental Marine Biology and Ecology
971:Journal of Experimental Marine Biology and Ecology
705:Bioturbation is important in the deep sea because
610:invertebrates to fish and marine mammals. In most
578:functions to increase the ease of fluid movement (
186:species are oriented with their heads towards the
4107:Palaeogeography, Palaeoclimatology, Palaeoecology
4104:
2649:Thibodeaux, Louis J.; Bierman, Victor J. (2003).
2648:
2323:Crowe, W. A.; Josefson, A. B.; Svane, I. (1987).
2078:Oceanography and Marine Biology: An Annual Review
1333:20.500.11755/e43f4d57-cf7f-494d-a724-a4b2aab2a772
905:Bioturbation's importance for soil processes and
792:appeared in the fossil record over a short time.
4154:
3355:
3271:
2739:
2374:
249:by other macrofaunal and microbial communities.
3115:Nitrogen Cycling in Coastal Marine Environments
2381:Proceedings of the National Academy of Sciences
1955:
4031:
3979:
3117:. John Wiley & Sons Ltd. pp. 301–338.
2789:
2199:Ortmann on migration of a mobile polychaete".
1425:
368:Bioturbation can either enhance or reduce the
1651:
1616:
16:Reworking of soils and sediments by organisms
3489:: CS1 maint: multiple names: authors list (
1132:
1130:
1128:
746:. Bioturbation is typically represented as D
4092:10.1306/212f7586-2b24-11d7-8648000102c1865d
2094:
865:rock. The study of such patterns is called
606:Major marine bioturbators range from small
3190:
462:, and thus contribute to the formation of
4013:
3961:
3904:
3746:
3615:
3438:
3381:
3248:
2674:
2533:
2474:
2418:
2400:
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1845:
1827:
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1734:
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1331:
1321:
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1190:
1125:
729:
333:
1708:
1444:
1390:
852:
645:
633:
503:
420:
236:
223:
18:
4143:Nereis Park (the World of Bioturbation)
2785:
2783:
2068:
1912:
363:
4155:
4073:
4071:
4027:
4025:
3975:
3973:
3833:
3831:
3829:
3665:
3663:
3452:
3450:
3128:
3126:
3124:
2841:
2839:
2837:
2781:
2779:
2777:
2775:
2773:
2771:
2769:
2767:
2765:
2763:
2735:
2733:
2731:
2729:
2655:Environmental Science & Technology
2603:Environmental Science & Technology
2556:Environmental Science & Technology
2492:
2490:
2488:
2486:
2440:
2438:
2370:
2368:
2279:
2277:
2194:
2112:
2110:
2103:(2): 150–178 – via Researchgate.
2064:
2062:
2060:
1908:
1906:
1756:
1754:
1593:
1591:
1589:
1587:
1585:
1583:
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1525:
1523:
1521:
1519:
1517:
1241:
1092:
1028:
1026:
1024:
1022:
1020:
877:. Thus, it affects the disciplines of
629:
3546:
3351:
3349:
3313:
3311:
3218:
3216:
3214:
3112:
3026:
3024:
3022:
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3018:
3016:
2996:
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2990:
2886:
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2880:
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2644:
2642:
2640:
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2549:
2547:
2545:
2496:
2036:
1479:
1291:
1289:
1287:
1237:
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1227:
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1212:
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1144:
1142:
1090:
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1080:
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1016:
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1010:
1008:
1006:
1004:
1002:
1000:
964:
962:
960:
958:
956:
954:
555:
244:keeping watch outside a shrimp burrow
3634:Estuarine, Coastal and Shelf Science
3003:American Fisheries Society Symposium
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405:
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3875:
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2037:Anker, Arthur; et al. (2005).
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1751:
1578:
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3456:
3346:
3308:
3211:
3013:
2987:
2875:
2637:
2590:
2542:
1284:
1220:
1207:
1139:
1069:
997:
951:
837:The evolution of trees during the
499:
54:. These include the alteration of
14:
4199:
4136:
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2849:Fundamental and Applied Limnology
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1035:Trends in Ecology & Evolution
788:, during which most major animal
346:begin to mix reduced sulfur from
287:lives within the deep portion of
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2861:10.1127/1863-9135/2007/0168-0027
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3798:
3763:
3698:
3672:Geochimica et Cosmochimica Acta
3624:
3583:
3540:
3497:
3406:
3265:
3184:
3161:
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2935:
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2316:
2238:) in Willapa Bay, Washington".
2227:
2188:
2133:
2088:
2030:
1984:
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528:(non-biting midges) larvae and
34:is defined as the reworking of
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3596:Marine Ecology Progress Series
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3274:Marine Ecology Progress Series
2329:Marine Ecology Progress Series
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1348:
1302:Marine Ecology Progress Series
1248:Marine Ecology Progress Series
1171:Marine Ecology Progress Series
873:, such as laminated layers or
744:partial differential equations
416:
1:
3569:10.1016/s0048-9697(02)00014-1
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3772:Journal of Marine Research
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3506:Limnology and Oceanography
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720:particulate organic carbon
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3526:10.4319/lo.1987.32.2.0313
3383:10.4319/lo.2009.54.6.2081
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3009:– via researchgate.
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2240:Marine Pollution Bulletin
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683:sulfate-reducing bacteria
601:
4054:10.1144/gsjgs.150.1.0141
1780:10.3389/fmars.2014.00041
1242:Pillay, D (2010-06-23).
622:, burrowing shrimp, and
188:sediment-water interface
61:Bioturbators are deemed
4163:Biological oceanography
3046:10.1139/cjfas-2014-0413
2402:10.1073/pnas.0902037106
2290:Journal of Sea Research
299:morphological evolution
278:symbiotic relationships
254:energetically favorable
3998:10.1098/rstb.1998.0195
3654:10.1006/ecss.1997.0298
3098:: CS1 maint: others (
2069:Karplus, Ilan (1987).
2022:: CS1 maint: others (
1978:10.1006/jare.1998.0477
1869:Wainright, SC (1990).
1570:: CS1 maint: others (
871:sedimentary structures
858:
803:Evolutionary Arms Race
730:Mathematical modelling
654:
643:
580:hydraulic conductivity
509:
466:Small mammals such as
426:
334:Biogeochemical effects
271:Macrotermes bellicosus
245:
234:
28:
4183:Physical oceanography
1679:10.1038/ismej.2009.62
1600:Symp. Zool. Soc. Lond
1099:Earth-Science Reviews
856:
652:marine nitrogen cycle
649:
637:
507:
425:Pocket gopher mounds
424:
240:
227:
22:
3170:Ecological Bulletins
2497:Boyle, R.A. (2014).
2197:Callianassa japonica
2043:Leuckart and Rüppel"
930:Argillipedoturbation
724:carbon sequestration
685:via the presence of
364:Organic contaminants
4119:2007PPP...245..382H
4046:1993JGSoc.150..141T
3954:2013Geo....41E.289B
3897:2011NatGe...4..372G
3853:2012Geo....40..395R
3723:2017NatSR...714287A
3684:2006GeCoA..70.5855R
3646:1998ECSS...46..531G
3608:2004MEPS..271....1S
3561:2002ScTEn.293..191B
3518:1987LimOc..32..313S
3431:1997DSRII..44.2295S
3425:(9–10): 2295–2317.
3374:2009LimOc..54.2081V
3332:1998DSRI...45..133A
3286:2016MEPS..557...31M
3241:2008CBio...18....1D
3147:1985JEMBE..85...75K
2957:2004FrBio..49..895M
2905:2011Ecol...92..373H
2804:2013AqInt..21....1A
2712:1996WatRe..30..704R
2667:2003EnST...37..252T
2615:2008EnST...42.1058G
2568:2010EnST...44.7456J
2518:2014NatGe...7..671B
2459:2016GeCoA.189..251D
2393:2009PNAS..106.8123C
2341:1987MEPS...41...61C
2302:1996JSR....36..217O
2252:2001MarPB..42..826D
2213:1988JEMBE.120...81T
2158:2007MarBi.152..611P
2142:Callianassa kraussi
1970:1999JArEn..41..123E
1927:1993FuEco...7....5H
1887:1990MEPS...62..271W
1820:2012PLoSO...744925C
1727:1972LimOc..17..583H
1670:2009ISMEJ...3.1269B
1631:1987JEMBE.107..219B
1498:1987MEPS...39...99P
1405:1984JEMBE..80...11W
1370:2006JEMBE.337..178M
1314:2010MEPS..399..173B
1260:2010MEPS..409..301P
1183:2012MEPS..446..285K
1111:2009ESRv...97..257W
1047:2006TEcoE..21..688M
983:2006JEMBE.330..403R
811:ecosystem functions
761:radioactive tracers
754:and, sometimes, an
630:Shallow and coastal
435:soil organic matter
150:such as clams, and
132:Nereis diversicolor
63:ecosystem engineers
3711:Scientific Reports
3617:10.3354/meps271001
2945:Freshwater Biology
2350:10.3354/meps041061
2047:Zoological Studies
1915:Functional Ecology
1896:10.3354/meps062271
1507:10.3354/meps039099
1480:Posey, MH (1987).
859:
786:Cambrian Explosion
707:deep-sea ecosystem
658:Coastal ecosystems
655:
644:
556:Rivers and streams
510:
508:Chironomid larvae.
427:
374:polluted sediments
246:
235:
138:Marenzelleria spp.
104:, mud shrimp, and
52:ecosystem services
29:
3992:(1365): 113–130.
3963:10.1130/g33606c.1
3885:Nature Geoscience
3678:(23): 5855–5867.
3294:10.3354/meps11837
2913:10.1890/09-1694.1
2676:10.1021/es032518j
2661:(13): 252A–258A.
2623:10.1021/es071607j
2576:10.1021/es100615g
2562:(19): 7456–7464.
2506:Nature Geoscience
2387:(20): 8123–8127.
2286:Pectinaria koreni
2236:Crassostrea gigas
1664:(11): 1269–1285.
1323:10.3354/meps08336
1269:10.3354/meps08628
1192:10.3354/meps09506
832:geological dating
827:Ediacaran Period.
679:nitrogen fixation
638:Bioturbation and
515:primary producers
444:Burrowing animals
406:Ecosystem impacts
184:Downward-conveyor
174:Arenicola marina,
122:Gallery-diffusers
112:Functional groups
4195:
4131:
4130:
4102:
4096:
4095:
4075:
4066:
4065:
4029:
4020:
4019:
4017:
3977:
3968:
3967:
3965:
3933:
3927:
3926:
3915:10.1038/ngeo1142
3908:
3879:
3873:
3872:
3861:10.1130/g32807.1
3835:
3824:
3823:
3802:
3796:
3795:
3767:
3761:
3760:
3750:
3702:
3696:
3695:
3667:
3658:
3657:
3628:
3622:
3621:
3619:
3587:
3581:
3580:
3555:(1–3): 191–200.
3544:
3538:
3537:
3501:
3495:
3494:
3488:
3480:
3454:
3445:
3444:
3442:
3410:
3404:
3403:
3385:
3368:(6): 2081–2088.
3353:
3344:
3343:
3315:
3306:
3305:
3269:
3263:
3262:
3252:
3220:
3209:
3208:
3188:
3182:
3181:
3165:
3159:
3158:
3130:
3119:
3118:
3110:
3104:
3103:
3097:
3089:
3064:
3058:
3057:
3040:(8): 1146–1158.
3028:
3011:
3010:
2998:
2985:
2984:
2939:
2933:
2932:
2888:
2873:
2872:
2843:
2832:
2831:
2787:
2758:
2757:
2737:
2724:
2723:
2695:
2689:
2688:
2678:
2646:
2635:
2634:
2609:(4): 1058–1065.
2597:
2588:
2587:
2551:
2540:
2539:
2537:
2526:10.1038/ngeo2213
2503:
2494:
2481:
2480:
2478:
2442:
2433:
2432:
2422:
2404:
2372:
2363:
2362:
2352:
2320:
2314:
2313:
2296:(3–4): 217–226.
2281:
2272:
2271:
2231:
2225:
2224:
2192:
2186:
2185:
2137:
2131:
2130:
2114:
2105:
2104:
2092:
2086:
2085:
2075:
2066:
2055:
2054:
2034:
2028:
2027:
2021:
2013:
1988:
1982:
1981:
1953:
1947:
1946:
1910:
1901:
1900:
1898:
1866:
1860:
1859:
1849:
1831:
1799:
1793:
1792:
1782:
1758:
1749:
1748:
1738:
1706:
1700:
1699:
1681:
1658:The ISME Journal
1649:
1643:
1642:
1614:
1608:
1607:
1595:
1576:
1575:
1569:
1561:
1535:
1512:
1511:
1509:
1477:
1471:
1470:
1442:
1436:
1435:
1423:
1417:
1416:
1388:
1382:
1381:
1352:
1346:
1345:
1335:
1325:
1293:
1282:
1281:
1271:
1239:
1218:
1214:
1205:
1204:
1194:
1162:
1137:
1134:
1123:
1122:
1094:
1067:
1066:
1030:
995:
994:
966:
917:Nathaniel Shaler
901:Research history
857:Planolite fossil
736:numerical models
612:marine sediments
493:biological pumps
411:Nutrient cycling
392:polychaete worms
357:(mineralization)
231:Arenicola marina
220:Ecological roles
162:Upward-conveyors
4203:
4202:
4198:
4197:
4196:
4194:
4193:
4192:
4168:Aquatic ecology
4153:
4152:
4139:
4134:
4103:
4099:
4076:
4069:
4030:
4023:
3978:
3971:
3934:
3930:
3906:10.1.1.717.5339
3880:
3876:
3836:
3827:
3803:
3799:
3768:
3764:
3703:
3699:
3668:
3661:
3629:
3625:
3588:
3584:
3545:
3541:
3502:
3498:
3482:
3481:
3469:
3455:
3448:
3411:
3407:
3354:
3347:
3316:
3309:
3270:
3266:
3229:Current Biology
3221:
3212:
3189:
3185:
3172:(35): 193–205.
3166:
3162:
3131:
3122:
3111:
3107:
3091:
3090:
3078:
3066:
3065:
3061:
3029:
3014:
2999:
2988:
2940:
2936:
2889:
2876:
2844:
2835:
2788:
2761:
2738:
2727:
2696:
2692:
2647:
2638:
2598:
2591:
2552:
2543:
2501:
2495:
2484:
2443:
2436:
2373:
2366:
2321:
2317:
2282:
2275:
2246:(10): 826–844.
2232:
2228:
2193:
2189:
2138:
2134:
2115:
2108:
2093:
2089:
2073:
2067:
2058:
2035:
2031:
2015:
2014:
2002:
1990:
1989:
1985:
1954:
1950:
1935:10.2307/2389861
1911:
1904:
1867:
1863:
1800:
1796:
1759:
1752:
1707:
1703:
1650:
1646:
1615:
1611:
1596:
1579:
1563:
1562:
1550:
1537:
1536:
1515:
1478:
1474:
1443:
1439:
1424:
1420:
1389:
1385:
1353:
1349:
1294:
1285:
1240:
1221:
1215:
1208:
1163:
1140:
1135:
1126:
1095:
1070:
1041:(12): 688–695.
1031:
998:
967:
952:
948:
926:
903:
851:
839:Devonian Period
782:
771:
749:
732:
703:
674:denitrification
632:
604:
558:
534:denitrification
530:tubificid worms
502:
500:Lakes and ponds
481:
419:
408:
384:, the invasive
366:
348:ocean sediments
340:ocean chemistry
336:
328:Deposit-feeding
222:
114:
17:
12:
11:
5:
4201:
4191:
4190:
4185:
4180:
4175:
4170:
4165:
4151:
4150:
4145:
4138:
4137:External links
4135:
4133:
4132:
4113:(3): 382–389.
4097:
4067:
4040:(1): 141–148.
4021:
3969:
3928:
3891:(6): 372–375.
3874:
3847:(5): 395–398.
3825:
3814:(2): 191–218.
3797:
3778:(1): 177–207.
3762:
3697:
3659:
3640:(4): 531–547.
3623:
3582:
3539:
3512:(2): 313–326.
3496:
3468:978-3642643996
3467:
3446:
3405:
3345:
3326:(1): 133–165.
3307:
3264:
3210:
3199:(2): 109–116.
3183:
3160:
3120:
3105:
3076:
3059:
3012:
2986:
2951:(7): 895–912.
2934:
2899:(2): 373–385.
2874:
2833:
2759:
2725:
2706:(3): 704–714.
2700:Water Research
2690:
2636:
2589:
2541:
2482:
2434:
2364:
2315:
2273:
2226:
2187:
2152:(3): 611–618.
2146:Marine Biology
2132:
2106:
2087:
2056:
2029:
2001:978-0226467283
2000:
1983:
1964:(2): 123–125.
1948:
1902:
1861:
1794:
1750:
1721:(4): 583–586.
1701:
1644:
1625:(3): 219–235.
1609:
1577:
1549:978-1570034312
1548:
1513:
1472:
1453:(4): 229–249.
1437:
1418:
1383:
1364:(2): 178–189.
1347:
1283:
1219:
1206:
1138:
1124:
1105:(1): 257–272.
1068:
996:
977:(1): 403–419.
949:
947:
944:
943:
942:
937:
932:
925:
922:
902:
899:
850:
847:
807:microbial mats
781:
778:
769:
747:
731:
728:
702:
699:
631:
628:
603:
600:
587:hyporheic zone
557:
554:
501:
498:
480:
477:
473:soil formation
468:pocket gophers
464:soil horizons.
460:soil biomantle
418:
415:
407:
404:
365:
362:
335:
332:
221:
218:
217:
216:
200:
199:
181:
159:
141:
113:
110:
90:pocket gophers
48:Charles Darwin
15:
9:
6:
4:
3:
2:
4200:
4189:
4188:Sedimentology
4186:
4184:
4181:
4179:
4176:
4174:
4171:
4169:
4166:
4164:
4161:
4160:
4158:
4149:
4146:
4144:
4141:
4140:
4128:
4124:
4120:
4116:
4112:
4108:
4101:
4093:
4089:
4085:
4081:
4074:
4072:
4063:
4059:
4055:
4051:
4047:
4043:
4039:
4035:
4028:
4026:
4016:
4011:
4007:
4003:
3999:
3995:
3991:
3987:
3983:
3976:
3974:
3964:
3959:
3955:
3951:
3947:
3943:
3939:
3932:
3924:
3920:
3916:
3912:
3907:
3902:
3898:
3894:
3890:
3886:
3878:
3870:
3866:
3862:
3858:
3854:
3850:
3846:
3842:
3834:
3832:
3830:
3821:
3817:
3813:
3809:
3801:
3793:
3789:
3785:
3781:
3777:
3773:
3766:
3758:
3754:
3749:
3744:
3740:
3736:
3732:
3728:
3724:
3720:
3716:
3712:
3708:
3701:
3693:
3689:
3685:
3681:
3677:
3673:
3666:
3664:
3655:
3651:
3647:
3643:
3639:
3635:
3627:
3618:
3613:
3609:
3605:
3601:
3597:
3593:
3586:
3578:
3574:
3570:
3566:
3562:
3558:
3554:
3550:
3543:
3535:
3531:
3527:
3523:
3519:
3515:
3511:
3507:
3500:
3492:
3486:
3478:
3474:
3470:
3464:
3460:
3453:
3451:
3441:
3436:
3432:
3428:
3424:
3420:
3416:
3409:
3401:
3397:
3393:
3389:
3384:
3379:
3375:
3371:
3367:
3363:
3359:
3352:
3350:
3341:
3337:
3333:
3329:
3325:
3321:
3314:
3312:
3303:
3299:
3295:
3291:
3287:
3283:
3279:
3275:
3268:
3260:
3256:
3251:
3246:
3242:
3238:
3234:
3230:
3226:
3219:
3217:
3215:
3206:
3202:
3198:
3194:
3187:
3179:
3175:
3171:
3164:
3156:
3152:
3148:
3144:
3140:
3136:
3129:
3127:
3125:
3116:
3109:
3101:
3095:
3087:
3083:
3079:
3077:9781118665442
3073:
3069:
3063:
3055:
3051:
3047:
3043:
3039:
3035:
3027:
3025:
3023:
3021:
3019:
3017:
3008:
3004:
2997:
2995:
2993:
2991:
2982:
2978:
2974:
2970:
2966:
2962:
2958:
2954:
2950:
2946:
2938:
2930:
2926:
2922:
2918:
2914:
2910:
2906:
2902:
2898:
2894:
2887:
2885:
2883:
2881:
2879:
2870:
2866:
2862:
2858:
2854:
2850:
2842:
2840:
2838:
2829:
2825:
2821:
2817:
2813:
2809:
2805:
2801:
2797:
2793:
2786:
2784:
2782:
2780:
2778:
2776:
2774:
2772:
2770:
2768:
2766:
2764:
2755:
2751:
2747:
2743:
2736:
2734:
2732:
2730:
2721:
2717:
2713:
2709:
2705:
2701:
2694:
2686:
2682:
2677:
2672:
2668:
2664:
2660:
2656:
2652:
2645:
2643:
2641:
2632:
2628:
2624:
2620:
2616:
2612:
2608:
2604:
2596:
2594:
2585:
2581:
2577:
2573:
2569:
2565:
2561:
2557:
2550:
2548:
2546:
2536:
2531:
2527:
2523:
2519:
2515:
2511:
2507:
2500:
2493:
2491:
2489:
2487:
2477:
2472:
2468:
2464:
2460:
2456:
2452:
2448:
2441:
2439:
2430:
2426:
2421:
2416:
2412:
2408:
2403:
2398:
2394:
2390:
2386:
2382:
2378:
2371:
2369:
2360:
2356:
2351:
2346:
2342:
2338:
2334:
2330:
2326:
2319:
2311:
2307:
2303:
2299:
2295:
2291:
2287:
2280:
2278:
2269:
2265:
2261:
2257:
2253:
2249:
2245:
2241:
2237:
2230:
2222:
2218:
2214:
2210:
2206:
2202:
2198:
2191:
2183:
2179:
2175:
2171:
2167:
2163:
2159:
2155:
2151:
2147:
2143:
2136:
2128:
2124:
2120:
2113:
2111:
2102:
2098:
2091:
2083:
2079:
2072:
2065:
2063:
2061:
2053:(2): 157–190.
2052:
2048:
2044:
2042:
2033:
2025:
2019:
2011:
2007:
2003:
1997:
1993:
1987:
1979:
1975:
1971:
1967:
1963:
1959:
1952:
1944:
1940:
1936:
1932:
1928:
1924:
1920:
1916:
1909:
1907:
1897:
1892:
1888:
1884:
1880:
1876:
1872:
1865:
1857:
1853:
1848:
1843:
1839:
1835:
1830:
1825:
1821:
1817:
1814:(9): e44925.
1813:
1809:
1805:
1798:
1790:
1786:
1781:
1776:
1772:
1768:
1764:
1757:
1755:
1746:
1742:
1737:
1732:
1728:
1724:
1720:
1716:
1712:
1705:
1697:
1693:
1689:
1685:
1680:
1675:
1671:
1667:
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1659:
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1624:
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1605:
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1594:
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1559:
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1528:
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1524:
1522:
1520:
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1487:
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1476:
1468:
1464:
1460:
1456:
1452:
1448:
1441:
1433:
1429:
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1414:
1410:
1406:
1402:
1398:
1394:
1387:
1379:
1375:
1371:
1367:
1363:
1359:
1351:
1343:
1339:
1334:
1329:
1324:
1319:
1315:
1311:
1307:
1303:
1299:
1292:
1290:
1288:
1279:
1275:
1270:
1265:
1261:
1257:
1253:
1249:
1245:
1238:
1236:
1234:
1232:
1230:
1228:
1226:
1224:
1213:
1211:
1202:
1198:
1193:
1188:
1184:
1180:
1176:
1172:
1168:
1161:
1159:
1157:
1155:
1153:
1151:
1149:
1147:
1145:
1143:
1133:
1131:
1129:
1120:
1116:
1112:
1108:
1104:
1100:
1093:
1091:
1089:
1087:
1085:
1083:
1081:
1079:
1077:
1075:
1073:
1064:
1060:
1056:
1052:
1048:
1044:
1040:
1036:
1029:
1027:
1025:
1023:
1021:
1019:
1017:
1015:
1013:
1011:
1009:
1007:
1005:
1003:
1001:
992:
988:
984:
980:
976:
972:
965:
963:
961:
959:
957:
955:
950:
941:
938:
936:
935:Bioirrigation
933:
931:
928:
927:
921:
918:
914:
913:
908:
907:geomorphology
898:
896:
892:
891:trace fossils
887:
884:
880:
879:sedimentology
876:
875:cross-bedding
872:
868:
864:
855:
849:Fossil record
846:
844:
840:
835:
833:
828:
824:
823:
819:
814:
812:
808:
804:
799:
795:
791:
787:
777:
773:
766:
765:radioisotopes
762:
757:
753:
745:
741:
737:
727:
725:
721:
716:
715:heterogeneity
712:
708:
698:
694:
692:
688:
684:
680:
675:
671:
670:nitrification
665:
663:
659:
653:
648:
641:
640:bioirrigation
636:
627:
625:
621:
617:
613:
609:
599:
596:
595:heterotrophic
592:
588:
583:
581:
577:
573:
569:
565:
562:
553:
550:
546:
541:
537:
535:
531:
527:
522:
518:
516:
506:
497:
494:
490:
486:
476:
474:
469:
465:
461:
458:known as the
457:
453:
452:soil porosity
449:
445:
441:
440:
436:
432:
423:
414:
412:
403:
401:
397:
393:
389:
388:
387:Marenzelleria
383:
379:
375:
371:
361:
358:
352:
349:
343:
341:
331:
329:
324:
320:
316:
312:
308:
302:
300:
296:
292:
291:
286:
285:
279:
276:evolution of
274:
272:
265:
262:
258:
255:
250:
243:
239:
233:
232:
226:
214:
210:
205:
202:
201:
197:
193:
189:
185:
182:
179:
175:
171:
167:
163:
160:
157:
153:
149:
145:
142:
140:
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134:
133:
128:
123:
120:
119:
118:
109:
107:
103:
99:
95:
91:
87:
83:
79:
77:
76:bioirrigation
73:
69:
68:trace fossils
64:
59:
57:
53:
49:
45:
41:
37:
33:
26:
21:
4110:
4106:
4100:
4083:
4079:
4037:
4033:
3989:
3985:
3945:
3941:
3931:
3888:
3884:
3877:
3844:
3840:
3811:
3807:
3800:
3775:
3771:
3765:
3717:(1): 14287.
3714:
3710:
3700:
3675:
3671:
3637:
3633:
3626:
3599:
3595:
3585:
3552:
3548:
3542:
3509:
3505:
3499:
3458:
3422:
3418:
3408:
3365:
3361:
3323:
3319:
3277:
3273:
3267:
3232:
3228:
3196:
3192:
3186:
3169:
3163:
3141:(1): 75–91.
3138:
3134:
3114:
3108:
3067:
3062:
3037:
3033:
3006:
3002:
2948:
2944:
2937:
2896:
2892:
2855:(1): 27–38.
2852:
2848:
2795:
2791:
2748:(1): 44–49.
2745:
2741:
2703:
2699:
2693:
2658:
2654:
2606:
2602:
2559:
2555:
2509:
2505:
2450:
2446:
2384:
2380:
2335:(1): 61–69.
2332:
2328:
2318:
2293:
2289:
2285:
2243:
2239:
2235:
2229:
2207:(1): 81–95.
2204:
2200:
2196:
2190:
2149:
2145:
2141:
2135:
2126:
2122:
2100:
2096:
2090:
2081:
2077:
2050:
2046:
2040:
2032:
1991:
1986:
1961:
1957:
1951:
1918:
1914:
1878:
1874:
1864:
1811:
1807:
1797:
1770:
1766:
1718:
1714:
1704:
1661:
1657:
1647:
1622:
1618:
1612:
1603:
1599:
1539:
1489:
1485:
1475:
1450:
1446:
1440:
1431:
1427:
1421:
1399:(1): 11–28.
1396:
1392:
1386:
1361:
1357:
1350:
1305:
1301:
1251:
1247:
1174:
1170:
1102:
1098:
1038:
1034:
974:
970:
910:
904:
888:
883:stratigraphy
860:
836:
820:
818:trace fossil
815:
783:
774:
763:such as Pb,
738:built using
733:
704:
695:
686:
666:
656:
605:
584:
576:salmon redds
566:
559:
542:
538:
523:
519:
511:
485:benthivorous
482:
442:
428:
409:
385:
367:
353:
344:
337:
303:
295:commensalist
288:
282:
268:
266:
251:
247:
229:
228:The lugworm
204:Regenerators
203:
196:Sipunculidae
183:
173:
161:
144:Biodiffusers
143:
136:
130:
121:
115:
106:midge larvae
102:ghost shrimp
80:
60:
44:biodiversity
32:Bioturbation
31:
30:
3948:(5): e289.
2798:(1): 1–17.
2535:10871/35799
2476:10871/23490
2453:: 251–268.
2129:(1): 21–34.
1921:(1): 5–12.
1881:: 271–281.
1308:: 173–186.
1254:: 301–303.
1177:: 285–302.
843:soil carbon
711:photic zone
691:nitrogenase
616:polychaetes
591:autotrophic
417:Terrestrial
400:hydrophobic
390:species of
323:recruitment
319:aquaculture
290:Callianassa
192:peanut worm
178:thalassinid
166:polychaetes
127:polychaetes
98:polychaetes
4157:Categories
3235:(1): 1–8.
2512:(9): 671.
2084:: 507–562.
1492:: 99–104.
946:References
889:Important
886:specimen.
526:chironomid
489:anadromous
479:Freshwater
382:Baltic Sea
307:gastropods
129:, such as
94:earthworms
72:diagenesis
4173:Limnology
4062:129182527
4006:0962-8436
3923:1752-0908
3901:CiteSeerX
3869:0091-7613
3792:129173448
3739:2045-2322
3534:1939-5590
3485:cite book
3477:851842693
3392:1939-5590
3302:0171-8630
3280:: 31–49.
3094:cite book
3086:798834896
3054:0706-652X
2973:1365-2427
2921:1939-9170
2820:0967-6120
2411:0027-8424
2174:0025-3162
2018:cite book
1838:1932-6203
1789:2296-7745
1745:1939-5590
1688:1751-7370
1566:cite book
1467:0036-4827
1342:0171-8630
1278:0171-8630
1201:0171-8630
940:Zoophycos
867:ichnology
863:lithified
822:Nenoxites
794:Predation
780:Evolution
756:advective
752:diffusion
693:) genes.
624:amphipods
549:turbidity
431:saprolite
315:amphipods
311:cumaceans
168:like the
156:epifaunal
56:nutrients
40:sediments
4178:Pedology
4148:Worm Cam
3757:29079758
3602:: 1–12.
3577:12109472
3400:53613556
3259:18164201
3178:20112854
2981:43255065
2929:21618917
2869:84815294
2828:15097632
2685:12875383
2631:18351072
2584:20831254
2429:19451639
2359:24827459
2268:11693637
2182:84904361
2010:43207081
1856:22984586
1808:PLOS ONE
1696:19458658
1558:47927758
1063:16901581
924:See also
740:ordinary
701:Deep sea
662:detritus
620:bivalves
608:infaunal
378:Invasive
257:oxidants
194:family,
180:shrimps.
152:amphipod
148:bivalves
82:Walruses
4115:Bibcode
4042:Bibcode
4015:1692181
3950:Bibcode
3942:Geology
3893:Bibcode
3849:Bibcode
3841:Geology
3748:5660215
3719:Bibcode
3680:Bibcode
3642:Bibcode
3604:Bibcode
3557:Bibcode
3514:Bibcode
3427:Bibcode
3370:Bibcode
3328:Bibcode
3282:Bibcode
3237:Bibcode
3205:4297030
3143:Bibcode
2953:Bibcode
2901:Bibcode
2893:Ecology
2800:Bibcode
2708:Bibcode
2663:Bibcode
2611:Bibcode
2564:Bibcode
2514:Bibcode
2455:Bibcode
2420:2688866
2389:Bibcode
2337:Bibcode
2298:Bibcode
2248:Bibcode
2209:Bibcode
2154:Bibcode
1966:Bibcode
1943:2389861
1923:Bibcode
1883:Bibcode
1847:3440345
1816:Bibcode
1723:Bibcode
1666:Bibcode
1627:Bibcode
1494:Bibcode
1401:Bibcode
1366:Bibcode
1310:Bibcode
1256:Bibcode
1179:Bibcode
1107:Bibcode
1043:Bibcode
979:Bibcode
798:armored
456:topsoil
396:infauna
209:fiddler
170:lugworm
4060:
4012:
4004:
3921:
3903:
3867:
3790:
3755:
3745:
3737:
3575:
3532:
3475:
3465:
3398:
3390:
3300:
3257:
3203:
3176:
3084:
3074:
3052:
2979:
2971:
2927:
2919:
2867:
2826:
2818:
2683:
2629:
2582:
2427:
2417:
2409:
2357:
2266:
2180:
2172:
2008:
1998:
1941:
1854:
1844:
1836:
1787:
1743:
1694:
1686:
1556:
1546:
1465:
1447:Sarsia
1340:
1276:
1217:66-80.
1199:
1061:
895:Eolian
602:Marine
568:Salmon
313:, and
242:Gobies
215:crabs.
88:, and
86:salmon
25:walrus
4086:(3).
4058:S2CID
3788:S2CID
3396:S2CID
3201:JSTOR
3174:JSTOR
2977:S2CID
2865:S2CID
2824:S2CID
2502:(PDF)
2355:JSTOR
2178:S2CID
2074:(PDF)
1939:JSTOR
790:phyla
572:redds
213:ghost
36:soils
4002:ISSN
3919:ISSN
3865:ISSN
3753:PMID
3735:ISSN
3573:PMID
3530:ISSN
3491:link
3473:OCLC
3463:ISBN
3388:ISSN
3298:ISSN
3255:PMID
3100:link
3082:OCLC
3072:ISBN
3050:ISSN
2969:ISSN
2925:PMID
2917:ISSN
2816:ISSN
2681:PMID
2627:PMID
2580:PMID
2425:PMID
2407:ISSN
2264:PMID
2170:ISSN
2024:link
2006:OCLC
1996:ISBN
1852:PMID
1834:ISSN
1785:ISSN
1741:ISSN
1692:PMID
1684:ISSN
1572:link
1554:OCLC
1544:ISBN
1463:ISSN
1338:ISSN
1274:ISSN
1197:ISSN
1059:PMID
881:and
742:and
650:The
545:carp
370:flux
211:and
176:and
135:and
38:and
4123:doi
4111:245
4088:doi
4050:doi
4038:150
4010:PMC
3994:doi
3990:353
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3911:doi
3857:doi
3816:doi
3780:doi
3743:PMC
3727:doi
3688:doi
3650:doi
3612:doi
3600:271
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3553:293
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3378:doi
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2853:168
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2716:doi
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2572:doi
2530:hdl
2522:doi
2471:hdl
2463:doi
2451:189
2415:PMC
2397:doi
2385:106
2345:doi
2306:doi
2256:doi
2217:doi
2205:120
2162:doi
2150:152
1974:doi
1931:doi
1891:doi
1842:PMC
1824:doi
1775:doi
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1635:doi
1623:107
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1362:337
1328:hdl
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770:B
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