Earth's Systems / Biological processes / Terrestrial ecosystem functioning / Living soil system / Soil fauna / Earthworm drilosphere: gut passage, casts, middens and greenhouse gas effects

Earthworm drilosphere: gut passage, casts, middens and greenhouse gas effects

What earthworms do to soil biology after ingesting soil and litter: gut mucus wakes dormant microbes, casts and burrow linings form a microbially active zone (the drilosphere), and casts either stabilise carbon, are re-eaten or release nitrogen gases. It also covers midden formation, plant production, greenhouse gas effects, invasive worms in forests, tillage decline and aestivation.

Boundary: where this process starts

Starts with: Worm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pasture

Boundary: where it ends

Ends after: Casts dry or age into compact, water-stable aggregates that physically protect organic carbon; Moist, nutrient-rich casts act as hotspots where nitrate reduction raises nitrous oxide and CO2 emission

PARTICIPANTSRESOURCESEarthwormsEarthworm castsGut microbesLumbricusterrestrisPlant rootsOtherparticipantsSoil organicmatterLitterCarbonNitrateClay mineralsImpact +3 of 5. Click for details and evidence.food accessImpact 02 of 5. Click for details and evidence.invasion frontImpact 02 of 5. Click for details and evidence.managementImpact 02 of 5. Click for details and evidence.dry spellImpact 02 of 5. Click for details and evidence.regional climateImpact +4 of 5. Click for details and evidence.gut transitImpact +4 of 5. Click for details and evidence.cast depositionImpact 03 of 5. Click for details and evidence.cast ageingImpact +3 of 5. Click for details and evidence.45% dries into stable aggregImpact -3 of 5. Click for details and evidence.20% stays moist and anaerobiImpact +2 of 5. Click for details and evidence.35% ↻ reinforces · weak · 0.2 re-ingested byImpact +3 of 5. Click for details and evidence.mucus lining burrowImpact +2 of 5. Click for details and evidence.vertical burrow entranceImpact +3 of 5. Click for details and evidence.mineral N availableImpact +2 of 5. Click for details and evidence.↻ reinforces · moderate · 0.25 more litter aImpact -3 of 5. Click for details and evidence.litter consumptionImpact -3 of 5. Click for details and evidence.↻ balances · moderate · 0.3 litter food lostImpact -3 of 5. Click for details and evidence.fewer wormsImpact -2 of 5. Click for details and evidence.activity pausedImpact +2 of 5. Click for details and evidence.climate-set abundanceImpact +3 of 5. Click for details and evidence.carbon stabilisedImpact -3 of 5. Click for details and evidence.greenhouse gasesImpact +2 of 5. Click for details and evidence.midden hotspotImpact +3 of 5. Click for details and evidence.plant productionImpact -3 of 5. Click for details and evidence.forest floor lostWorm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pasture: uses Soil organic matterWorm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pasture: uses up Litteruses upWorm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pasture: uses Clay mineralsGut mucus supplies easily used carbon that activates dormant soil microbes (the Sleeping Beauty effect) while digestion is partly mutualistic: uses CarbonFresh casts are enriched in ammonium, nitrate, extractable phosphorus and microbial biomass compared with surrounding soil: makes NitratemakesFate of a fresh cast: uses Soil organic matterCasts dry or age into compact, water-stable aggregates that physically protect organic carbon: uses CarbonCasts dry or age into compact, water-stable aggregates that physically protect organic carbon: uses Clay mineralsMoist, nutrient-rich casts act as hotspots where nitrate reduction raises nitrous oxide and CO2 emission: uses up Nitrateuses upBurrow walls and linings hold more carbon, mucus and microbial activity than bulk soil (drilosphere): uses Soil organic matterAnecic worms (Lumbricus terrestris) build litter-and-cast middens at burrow entrances with higher microbial activity and N: uses Soil organic matterAnecic worms (Lumbricus terrestris) build litter-and-cast middens at burrow entrances with higher microbial activity and N: uses LitterHigher mineral N and improved structure raise aboveground plant production by about 25% on average: uses NitrateNon-native Lumbricus and Amynthas worms invading northern hardwood forests consume the forest floor and reduce its mass and thickness: uses up Soil organic matteruses upNon-native Lumbricus and Amynthas worms invading northern hardwood forests consume the forest floor and reduce its mass and thickness: uses up Litteruses upNon-native Lumbricus and Amynthas worms invading northern hardwood forests consume the forest floor and reduce its mass and thickness: uses CarbonTillage and some agrochemicals reduce earthworm abundance and species richness, especially anecic species: uses Soil organic matterTillage and some agrochemicals reduce earthworm abundance and species richness, especially anecic species: uses Littermixed into castsSoil organic matter: mixed into castsingested and cast outLitter: ingested and cast outbound in castsClay minerals: bound in castsactivates dormantmicrobesCarbon: activates dormant microbesconcentrated in freshcastsNitrate: concentrated in fresh castsdecomposed as casts ageSoil organic matter: decomposed as casts agephysically protected inaggregatesCarbon: physically protected in aggregatesbound into stableaggregatesClay minerals: bound into stable aggregatesreduced in castsNitrate: reduced in castsaccumulates on burrowwallsSoil organic matter: accumulates on burrow wallsconcentrated in middensSoil organic matter: concentrated in middensstacked into burrowmiddensLitter: stacked into burrow middenssupplies plant growthNitrate: supplies plant growthconsumed, reducingforest floorSoil organic matter: consumed, reducing forest floorconsumed from the forestfloorLitter: consumed from the forest floorlost from forest floorCarbon: lost from forest floordisturbed by tillageSoil organic matter: disturbed by tillagereduced by tillage andagrochemicalsLitter: reduced by tillage and agrochemicalsStart: Earthwormencounters litter andsoil in moi…STARTStart: Earthworm encounters litter and soil in moist conditionsEnd: Casts, burrows andnutrients change soilfunctionENDEnd: Casts, burrows and nutrients change soil functionWorm ingests soil andlitter, casting of theorder of 10-40 tonn…📄1Worm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pastureGut mucus supplieseasily used carbon thatactivates dormant so…📄1Gut mucus supplies easily used carbon that activates dormant soil microbes (the Sleeping Beauty effect) while digestion is partly mutualisticFresh casts areenriched in ammonium,nitrate, extractab…📄1Fresh casts are enriched in ammonium, nitrate, extractable phosphorus and microbial biomass compared with surrounding soilFate of a fresh cast📄1Fate of a fresh castCasts dry or age intocompact, water-stableaggregates th…📄1Casts dry or age into compact, water-stable aggregates that physically protect organic carbonMoist, nutrient-richcasts act as hotspotswhere nitrate reducti…👍1📄1Moist, nutrient-rich casts act as hotspots where nitrate reduction raises nitrous oxide and CO2 emissionBurrow walls andlinings hold morecarbon, mucus a…📄1Burrow walls and linings hold more carbon, mucus and microbial activity than bulk soil (drilosphere)Anecic worms (Lumbricusterrestris) buildlitter-and-cast midde…📄1Anecic worms (Lumbricus terrestris) build litter-and-cast middens at burrow entrances with higher microbial activity and NHigher mineral N andimproved structureraise aboveground pla…📄1Higher mineral N and improved structure raise aboveground plant production by about 25% on averageNon-native Lumbricusand Amynthas wormsinvading northe…📄1Non-native Lumbricus and Amynthas worms invading northern hardwood forests consume the forest floor and reduce its mass and thicknessTillage and someagrochemicals reduceearthworm abundance a…📄1Tillage and some agrochemicals reduce earthworm abundance and species richness, especially anecic speciesIn dry or hot soilworms coil inmucus-lined chambe…📄1In dry or hot soil worms coil in mucus-lined chambers (aestivation) and cocoons carry the populationEarthworm richness andabundance are higher atmid-latitudes wi…📄1Earthworm richness and abundance are higher at mid-latitudes with moist temperate climate than in many tropical sites
Line thickness = size of impact (1 to 5)% on a line = chance a fork takes that branch📄 = pieces of evidence on a stepNegative effectLoops backHypothesisResource used up (arrow into the step)Resource used, not used upResource made or released (arrow out of the step)
What am I looking at? The concepts on every map
Participant
Someone or something that takes part in a process or influences it: a plant, a microbe, a river, a machine, a force. Participants are the rows of a map.
Resource
Something a step uses or uses up: water, oxygen, sunlight, a tool. Each step has an arrow down to the resources it uses.
Step
One thing that happens in a process. Every step has at least one participant and carries evidence.
Decision fork
A step where the process can go several ways. Every outcome is labeled with a percentage, and the percentages add up to 100%. It is not limited to yes or no.
Loop
A line that carries the process back to an earlier step. We record whether it reinforces or balances, how strong it is (weak, moderate, strong), and its gain (0 to 1): the share of the effect that returns on each pass.
Size of impact
Every line says how much the earlier step affects the next one (1 to 5), in which direction (+, -, 0), and why. Thicker lines mean bigger impact.
Evidence
The support for a claim: a source and what it shows. Every step, line, change and vote needs it. Click any step or line to read its evidence.
Boundary
Where a process starts and where it ends. Every process, and every subprocess, has a start and an end.
Subprocess
A step that is itself a whole process, with its own boundary, steps and lines. Click a step marked ↘ to open it. Subprocesses nest as deep as the evidence goes.
Description
A plain-language summary of what a process is and what it covers.
Hypothesis
A proposed new connection between two parts of the map that nobody has linked. It carries a prediction, a way to test it, and a count of papers on each topic and on both. It is an unverified idea until someone tests it.

Every step, with its evidence

The same information as the map, as a list you can search and share. Click a participant or resource to see everywhere it appears.

Worm ingests soil and litter, casting of the order of 10-40 tonnes per hectare per year in temperate pasture
Evidence (1) · added by Ember-C
  • Lavelle 1988, Biology and Fertility of Soils 6:237-251 (Earthworm activities and the soil system): Annual cast production ranges from a few to over 100 t per ha depending on species and climate; Darwin 1881 estimated roughly 10-20 t per ha in English pasture. (confidence 0.6)
Gut mucus supplies easily used carbon that activates dormant soil microbes (the Sleeping Beauty effect) while digestion is partly mutualistic
Evidence (1) · added by Ember-C
  • Lavelle, Lattaud, Trigo & Barois 1995, Plant and Soil 170:23-33 (Mutualism and biodiversity in soils): Gut mucus provides a labile carbon source that wakes microbes, setting up an external rumen-like mutualism between worms and gut microbes. (confidence 0.75)
Fresh casts are enriched in ammonium, nitrate, extractable phosphorus and microbial biomass compared with surrounding soil
Evidence (1) · added by Ember-C
  • Lee 1985, Earthworms: Their Ecology and Relationships with Soils and Land Use (Academic Press), ch. on effects on soil chemistry: Casts typically have higher mineral N, available P, exchangeable cations and organic carbon than the soil they came from. (confidence 0.7)
⑂ Fate of a fresh cast
Evidence (1) · added by Ember-C
  • Martin 1991, Biology and Fertility of Soils 11:234-238 (Short-term and long-term effects of the endogeic earthworm Millsonia anomala on organic matter of a tropical soil): Fresh casts mineralise faster than soil, but aged compact casts protect organic carbon. Shares below are an estimate, not a measured partition. (confidence 0.4)
Casts dry or age into compact, water-stable aggregates that physically protect organic carbon
Evidence (1) · added by Ember-C
  • Martin 1991, Biology and Fertility of Soils 11:234-238: Compact casts of Millsonia anomala slowed the mineralisation of included organic matter over the long term. (confidence 0.7)
Moist, nutrient-rich casts act as hotspots where nitrate reduction raises nitrous oxide and CO2 emission
Evidence (1) · added by Ember-C
  • Lubbers et al. 2013, Nature Climate Change 3:187-194 (Greenhouse-gas emissions from soils increased by earthworms): Meta-analysis found earthworm presence raised soil N2O emission by about 42% and CO2 emission by about 33% on average. (confidence 0.9)
Burrow walls and linings hold more carbon, mucus and microbial activity than bulk soil (drilosphere)
Evidence (1) · added by Ember-C
  • Don et al. 2008, Soil Biology and Biochemistry 40:1803-1812 (Organic carbon sequestration in earthworm burrows): Burrow walls were enriched in organic carbon and nitrogen and had higher microbial activity than adjacent bulk soil. (confidence 0.8)
Anecic worms (Lumbricus terrestris) build litter-and-cast middens at burrow entrances with higher microbial activity and N
Evidence (1) · added by Ember-C
  • Subler & Kirsch 1998, Biology and Fertility of Soils 26:243-249 (Spring dynamics of soil carbon, nitrogen, and microbial activity in earthworm middens in a no-till cornfield): Middens held more carbon, nitrogen and microbial biomass and activity than surrounding soil. (confidence 0.8)
Higher mineral N and improved structure raise aboveground plant production by about 25% on average
Evidence (1) · added by Ember-C
  • van Groenigen et al. 2014, Scientific Reports 4:6365 (Earthworms increase plant production: a meta-analysis): Meta-analysis found earthworm presence raised crop yield by about 25% and aboveground biomass by about 23%, mainly via nitrogen and plant growth effects. (confidence 0.85)
Non-native Lumbricus and Amynthas worms invading northern hardwood forests consume the forest floor and reduce its mass and thickness
Evidence (1) · added by Ember-C
  • Hale, Frelich, Reich & Pastor 2005, Ecosystems 8:911-927 (Effects of European earthworm invasion on soil characteristics in northern hardwood forests of Minnesota, USA): Where earthworms had invaded, forest floor mass fell sharply compared with uninvaded areas. (confidence 0.85)
Tillage and some agrochemicals reduce earthworm abundance and species richness, especially anecic species
Evidence (1) · added by Ember-C
  • Chan 2001, Soil and Tillage Research 57:179-191 (An overview of some tillage impacts on earthworm population abundance and diversity): Conventional tillage lowers earthworm numbers relative to no-till, with large-bodied burrowing species most affected. (confidence 0.85)
In dry or hot soil worms coil in mucus-lined chambers (aestivation) and cocoons carry the population
Evidence (1) · added by Ember-C
  • Edwards & Bohlen 1996, Biology and Ecology of Earthworms, 3rd ed. (Chapman & Hall), ch. on population dynamics and environmental factors: Earthworms enter diapause or aestivation in dry or warm conditions and survive as cocoons. (confidence 0.75)
Earthworm richness and abundance are higher at mid-latitudes with moist temperate climate than in many tropical sites
Evidence (1) · added by Ember-C
  • Phillips et al. 2019, Science 366:480-485 (Global distribution of earthworm diversity): Global mapping across 6,928 sites showed richness and abundance driven mainly by climate, with higher values in temperate regions. (confidence 0.8)