Biomass (ecology)

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Biomass (ecology)
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{{Use dmy dates|date=June 2013}}{{multiple image
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| caption1 = Apart from bacteria, the total global live biomass has been estimated as 550 or 560 billion tonnes C, most of which is found in forests.WEB,weblink Biomass, yes,weblink" title="">weblink 14 June 2010, dmy-all,
| image2 = Klamath river estuary.jpg
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| caption2 = Shallow aquatic environments, such as wetlands, estuaries and coral reefs, can be as productive as forests, generating similar amounts of new biomass each year on a given area.
}}The biomass is the mass of living biological organisms in a given area or ecosystem at a given time. Biomass can refer to species biomass, which is the mass of one or more species, or to community biomass, which is the mass of all species in the community. It can include microorganisms, plants or animals.{{GoldBookRef|title=biomass|url=}} The mass can be expressed as the average mass per unit area, or as the total mass in the community.How biomass is measured depends on why it is being measured. Sometimes, the biomass is regarded as the natural mass of organisms in situ, just as they are. For example, in a salmon fishery, the salmon biomass might be regarded as the total wet weight the salmon would have if they were taken out of the water. In other contexts, biomass can be measured in terms of the dried organic mass, so perhaps only 30% of the actual weight might count, the rest being water. For other purposes, only biological tissues count, and teeth, bones and shells are excluded. In some applications, biomass is measured as the mass of organically bound carbon (C) that is present.The total live biomass on Earth is about 550–560 billion tonnes C,JOURNAL, Bar-On YM, Phillips R, Milo R, The biomass distribution on Earth, Proceedings of the National Academy of Sciences of the United States of America, 115, 25, 6506–6511, June 2018, 29784790, 6016768, 10.1073/pnas.1711842115,weblink 1998PNAS...95.6578W, and the total annual primary production of biomass is just over 100 billion tonnes C/yr. The total live biomass of bacteria may be as much as that of plants and animals or may be much less. The total number of DNA base pairs on Earth, as a possible approximation of global biodiversity, is estimated at (5.3±3.6){{e|37}}, and weighs 50 billion tonnes.JOURNAL, Landenmark HK, Forgan DH, Cockell CS, An Estimate of the Total DNA in the Biosphere, PLoS Biology, 13, 6, e1002168, June 2015, 26066900, 4466264, 10.1371/journal.pbio.1002168, NEWS, Nuwer, Rachel, vanc, 18 July 2015, Counting All the DNA on Earth,weblink The New York Times, New York, The New York Times Company, 0362-4331, 2015-07-18, In comparison, the total mass of the biosphere has been estimated to be as much as 4{{e|12}} tonnes of carbon.WEB,weblink The Biosphere: Diversity of Life, Aspen Global Change Institute, Basalt, CO, 2015-07-19,

Ecological pyramids

(File:Ecological pyramid.svg|thumb|An ecological pyramid.)An ecological pyramid is a graphical representation that shows, for a given ecosystem, the relationship between biomass or biological productivity and trophic levels.
  • A biomass pyramid shows the amount of biomass at each trophic level.
  • A productivity pyramid shows the production or turn-over in biomass at each trophic level.
An ecological pyramid provides a snapshot in time of an ecological community.The bottom of the pyramid represents the primary producers (autotrophs). The primary producers take energy from the environment in the form of sunlight or inorganic chemicals and use it to create energy-rich molecules such as carbohydrates. This mechanism is called primary production. The pyramid then proceeds through the various trophic levels to the apex predators at the top.When energy is transferred from one trophic level to the next, typically only ten percent is used to build new biomass. The remaining ninety percent goes to metabolic processes or is dissipated as heat. This energy loss means that productivity pyramids are never inverted, and generally limits food chains to about six levels. However, in oceans, biomass pyramids can be wholly or partially inverted, with more biomass at higher levels.{{clear}}

Terrestrial biomass

Terrestrial biomass generally decreases markedly at each higher trophic level (plants, herbivores, carnivores). Examples of terrestrial producers are grasses, trees and shrubs. These have a much higher biomass than the animals that consume them, such as deer, zebras and insects. The level with the least biomass are the highest predators in the food chain, such as foxes and eagles.In a temperate grassland, grasses and other plants are the primary producers at the bottom of the pyramid. Then come the primary consumers, such as grasshoppers, voles and bison, followed by the secondary consumers, shrews, hawks and small cats. Finally the tertiary consumers, large cats and wolves. The biomass pyramid decreases markedly at each higher level.

Ocean biomass

{{marine food chain}}{{see also|Marine life}}Ocean or marine biomass, in a reversal of terrestrial biomass, can increase at higher trophic levels. In the ocean, the food chain typically starts with phytoplankton, and follows the course:Phytoplankton → zooplankton → predatory zooplankton → filter feeders → predatory fishPhytoplankton are the main primary producers at the bottom of the marine food chain. Phytoplankton use photosynthesis to convert inorganic carbon into protoplasm. They are then consumed by microscopic animals called zooplankton.Zooplankton comprise the second level in the food chain, and includes small crustaceans, such as copepods and krill, and the larva of fish, squid, lobsters and crabs.In turn, small zooplankton are consumed by both larger predatory zooplankters, such as krill, and by forage fish, which are small, schooling, filter-feeding fish. This makes up the third level in the food chain.{{clear}}(File:Arctic food web.svg|thumb|300px|left|An ocean food web showing a network of food chains)The fourth trophic level consists of predatory fish, marine mammals and seabirds that consume forage fish. Examples are swordfish, seals and gannets.Apex predators, such as orcas, which can consume seals, and shortfin mako sharks, which can consume swordfish, make up the fifth trophic level. Baleen whales can consume zooplankton and krill directly, leading to a food chain with only three or four trophic levels.Marine environments can have inverted biomass pyramids. In particular, the biomass of consumers (copepods, krill, shrimp, forage fish) is larger than the biomass of primary producers. This happens because the ocean's primary producers are tiny phytoplankton that grow and reproduce rapidly, so a small mass can have a fast rate of primary production. In contrast, terrestrial primary producers grow and reproduce slowly.{{clear}}There is an exception with cyanobacteria. Marine cyanobacteria are the smallest known photosynthetic organisms; the smallest of all, Prochlorococcus, is just 0.5 to 0.8 micrometres across.JOURNAL, Kettler GC, Martiny AC, Huang K, Zucker J, Coleman ML, Rodrigue S, Chen F, Lapidus A, Ferriera S, Johnson J, Steglich C, Church GM, Richardson P, Chisholm SW, Patterns and implications of gene gain and loss in the evolution of Prochlorococcus, PLoS Genetics, 3, 12, e231, December 2007, 18159947, 2151091, 10.1371/journal.pgen.0030231, Prochlorococcus is possibly the most plentiful species on Earth: a single millilitre of surface seawater may contain 100,000 cells or more. Worldwide, there are estimated to be several octillion (~1027) individuals.APOD, 27 September 2006, Earth from Saturn, Prochlorococcus is ubiquitous between 40°N and 40°S and dominates in the oligotrophic (nutrient poor) regions of the oceans.JOURNAL, Partensky F, Hess WR, Vaulot D, Prochlorococcus, a marine photosynthetic prokaryote of global significance, Microbiology and Molecular Biology Reviews, 63, 1, 106–27, March 1999, 10066832, 98958, The bacterium accounts for an estimated 20% of the oxygen in the Earth's atmosphere, and forms part of the base of the ocean food chain.WEB,weblink The Most Important Microbe You've Never Heard Of,,

Bacterial biomass

There are typically 50 million bacterial cells in a gram of soil and a million bacterial cells in a millilitre of fresh water. In a much-cited study from 1998,JOURNAL, Whitman WB, Coleman DC, Wiebe WJ, Prokaryotes: the unseen majority, Proceedings of the National Academy of Sciences of the United States of America, 95, 12, 6578–83, June 1998, 9618454, 33863, 10.1073/pnas.95.12.6578,weblink 1998PNAS...95.6578W, the world bacterial biomass had been mistakenly calculated to be 350 to 550 billions of tonnes of carbon, equal to between 60% and 100% of the carbon in plants. More recent studies of seafloor microbes cast considerable doubt on that; one study in 2012JOURNAL, Kallmeyer J, Pockalny R, Adhikari RR, Smith DC, D'Hondt S, Global distribution of microbial abundance and biomass in subseafloor sediment, Proceedings of the National Academy of Sciences of the United States of America, 109, 40, 16213–6, October 2012, 22927371, 3479597, 10.1073/pnas.1203849109,weblink 2012PNAS..10916213K, reduced the calculated microbial biomass on the seafloor from the original 303 billions of tonnes of C to just 4.1 billions of tonnes of C, reducing the global biomass of prokaryotes to 50 to 250 billions of tonnes of C. Further, if the average per-cell biomass of prokaryotes is reduced from 86 to 14 femtograms C, then the global biomass of prokaryotes was reduced to 13 to 44.5 billions of tonnes of C, equal to between 2.4% and 8.1% of the carbon in plants. As of 2018, there continues to be some controversy over what the global bacterial biomass is. A census published by the PNAS in May 2018 gives for bacterial biomass ~70 billions of tonnes of carbon, equal to 15% of the whole biomass. A census by the Deep Carbon Observatory project published in December 2018 gives a smaller figure of up to 23 billion tonnes of carbon.NEWS, Deep Carbon Observatory, Life in deep Earth totals 15 to 23 billion tons of carbon -- hundreds of times more than humans - Deep Carbon Observatory collaborators, exploring the 'Galapagos of the deep,' add to what's known, unknown, and unknowable about Earth's most pristine ecosystem,weblink 10 December 2018, EurekAlert!, 11 December 2018, NEWS, Dockrill, Peter, Scientists Reveal a Massive Biosphere of Life Hidden Under Earth's Surface,weblink 11 December 2018, Science Alert, 11 December 2018, NEWS, Gabbatiss, Josh, Massive ‘deep life’ study reveals billions of tonnes of microbes living far beneath Earth’s surface,weblink 11 December 2018, The Independent, 11 December 2018, {| class=wikitable!Geographic location!Number of cells (× 10{{sup|29}})!Billions of tonnes of carbon
Open ocean}}1.2}}1.7 to 10}}
Terrestrial soil}}2.6}}3.7 to 22}}
Subsurface terrestrial}}2.5 to 25}}3.5 to 215}}

Global biomass

Estimates for the global biomass of species and higher level groups are not always consistent across the literature. The total global biomass has been estimated at about 550 billion tonnes C.Groombridge B, Jenkins MD (2000) Global biodiversity: Earth’s living resources in the 21st century Page 11. World Conservation Monitoring Centre, World Conservation Press, CambridgeJOURNAL, Bar-On, Yinon M., Phillips, Rob, Milo, Ron, 2018-06-19, The biomass distribution on Earth,weblink Proceedings of the National Academy of Sciences, en, 115, 25, 6506–6511, 10.1073/pnas.1711842115, 0027-8424, 29784790, Most of this biomass is found on land, with only 5 to 10 billion tonnes C found in the oceans. On land, there is about 1,000 times more plant biomass (phytomass) than animal biomass (zoomass). About 18% of this plant biomass is eaten by the land animals.Hartley, Sue (2010) The 300 Million Years War: Plant Biomass v Herbivores Royal Institution Christmas Lecture. However, in the ocean, the animal biomass is nearly 30 times larger than the plant biomass.Darlington, P (1966)weblink "Biogeografia". Published in The Great Soviet Encyclopedia, 3rd Edition (1970–1979). Most ocean plant biomass is eaten by the ocean animals.{| class=wikitable!!name!number of species!date of estimate!individual count!mean living mass of individual!percent biomass (dried)!total number of carbon atoms!global dry biomass in million tonnes!global wet (fresh) biomass in million tonnes! rowspan=8 |Terrestrial
{{center|Humans}} {{center|1}}2012US world population clock {{webarchive weblink >date=1 July 2009 }}}}7.0 billion}}50 kg(incl children)}}30%}}4.015{{e|36}}Freitas, Robert A. Jr.Nanomedicine 3.1 Human Body Chemical Composition Foresight Institute, 1998}}105}}350}}
2005}}4.63 billion}}62 kg(excl. children)}}|||287WALPOLE SC, PRIETO-MERINO D, EDWARDS P, CLELAND J, STEVENS G, ROBERTS I > TITLE = THE WEIGHT OF NATIONS: AN ESTIMATION OF ADULT HUMAN BIOMASS VOLUME = 12 PAGES = 439 PMID = 22709383 DOI = 10.1186/1471-2458-12-439 POSTSCRIPT =, }}
Cattle}}1}}|1.3 billionCATTLE TODAY >URL=HTTP://CATTLE-TODAY.COM/ PUBLISHER=CATTLE-TODAY.COM, 2013-10-15, }}400 kg}}30%}}|156}}520}}
Sheep and goats}}2}}2002}}1.75 billionWorld's Rangelands Deteriorating Under Mounting Pressure {{webarchiveweblink >date=11 March 2008 }} Earth Policy Institute 2002}}60 kg}}30%}}|31.5}}105}}
Chickens}}1}}|24 billion}}2 kg}}30%}}|14.4}}48}}
Ants}}12,64weblink}}|107–108 billionEMBERY >FIRST1=JOAN FIRST2=ED FIRST3=HAVLICEK TITLE=JOAN EMBERY'S COLLECTION OF AMAZING ANIMAL FACTS PUBLISHER=DELACORTE PRESS ISBN=978-0-385-28486-8, }}3{{e|−6}} kg (0.003 grams)}}30%}}|10–100}}30-300}}
Earthworms}}>7,000}}1881Darwin}}1.3{{e URL = HTTPS://VERMECOLOGY.WORDPRESS.COM/2017/02/12/NATURE-ARTICLE-TO-COMMEMORATE-CHARLES-DARWINS-BIRTHDAY-ON-12TH-FEB/, Darwin's win-win for Global Worming?, }}3 g}}30% LEE KE >TITLE=EARTHWORMS: THEIR ECOLOGY AND RELATIONSHIPS WITH SOILS AND LAND USE PUBLISHER=ACADEMIC PRESS ISBN=978-0-12-440860-9, }}|1,140–2,280}}3,800–7,600}}
Termites}}>2,800}}1996}}|||||445Sum of [(biomass m{{sup2}})] from table 3 in Sanderson, M.G. 1996 Biomass of termites and their emissions of methane and carbon dioxide: A global database Global Biochemical Cycles, Vol 10:4 543-557}}
! rowspan=6 | {{center|Marine}}
Fish}}>10,000}}2009}}||||| 800-2,000Wilson RW, Millero FJ, Taylor JR, Walsh PJ, Christensen V, Jennings S and Grosell M (2009) "Contribution of Fish to the Marine Inorganic Carbon Cycle" Science, 323 (5912) 359–362. (This article provides a first estimate of global fish "wet weight" biomass)}}
Antarctic krill}}1}}1924–2004}}7.8{{e|14}}}}0.486 g}}|||379}}
Copepods(a zooplankton)}} 13,000}}||10−6–10−9 kg }}|1{{e TITLE = BIOGEOCHEMICAL FLUXES THROUGH MESOZOOPLANKTON JOURNAL = GLOBAL BIOGEOCHEMICAL CYCLES ISSUE = 2 DOI = 10.1029/2005GB002511, 2006GBioC..20.2003B, }}||
! rowspan=1 |Global
Prokaryotes(bacteria)}}?}}2018}}1{{e|31}} cells}}||| 23,000 – 70,000}}|
Humans comprise about 100 million tonnes of the Earth's dry biomass,The world human population was 6.6 billion in January 2008. At an average weight of 100 pounds (30 lbs of biomass), that equals 100 million tonnes.{{clarify| For this value, more than 7bn people are needed in my calculation. And the wet biomass "420Mt" claim doesn't fit to this either|date=April 2012}} domesticated animals about 700 million tonnes, earthworms over 1,100 million tonnes, and annual cereal crops about 2.3 billion tonnes.FAO Statistical Yearbook 2013: page 130 -weblink most successful animal species, in terms of biomass, may well be Antarctic krill, Euphausia superba, with a fresh biomass approaching 500 million tonnes,JOURNAL, 10.1016/j.dsr.2008.12.007, Atkinson A, Siegel V, Pakhomov EA, Jessopp MJ, Loeb V, 2009, A re-appraisal of the total biomass and annual production of Antarctic krill, Deep-Sea Research Part I, 56, 727–740,weblink 5, 2009DSRI...56..727A, Ross, R. M. and Quetin, L. B. (1988). Euphausia superba: a critical review of annual production. Comp. Biochem. Physiol. 90B, 499-505. although domestic cattle may also reach these immense figures.{{Citation needed|date=September 2011}} However, as a group, the small aquatic crustaceans called copepods may form the largest animal biomass on earth.WEB,weblink Biology of Copepods,weblink" title="">weblink 1 January 2009, Carl von Ossietzky University of Oldenburg,, A 2009 paper in Science estimates, for the first time, the total world fish biomass as somewhere between 0.8 and 2.0 billion tonnes.JOURNAL, Wilson RW, Millero FJ, Taylor JR, Walsh PJ, Christensen V, Jennings S, Grosell M, Contribution of fish to the marine inorganic carbon cycle, Science, 323, 5912, 359–362, January 2009, 19150840, 10.1126/science.1157972,weblink 2009Sci...323..359W, Researcher gives first-ever estimate of worldwide fish biomass and impact on climate change, 15 January 2009. It has been estimated that about 1% of the global biomass is due to phytoplankton,JOURNAL, Bidle KD, Falkowski PG, Cell death in planktonic, photosynthetic microorganisms, Nature Reviews. Microbiology, 2, 8, 643–655, August 2004, 15263899, 10.1038/nrmicro956,weblink and 25% is due to fungi.JOURNAL, Miller, JD, 1992, Fungi as contaminants in indoor air,weblink Atmospheric Environment, 26, 12, 2163–2172, 10.1016/0960-1686(92)90404-9, 1992AtmEn..26.2163M, JOURNAL, Sorenson WG, Fungal spores: hazardous to health?, Environmental Health Perspectives, 107 Suppl 3, Suppl 3, 469–472, June 1999, 10423389, 1566211, 10.1289/ehp.99107s3469, File:Cytisus scoparius2.jpg|Grasses, trees and shrubs have a much higher biomass than the animals that consume themFile:Bluegreen algae.jpg|The total biomass of bacteria may equal that of plants.File:copepodkils.jpg|Copepods may form the largest biomass of any animal species group.File:Antarctic krill (Euphausia superba).jpg|Antarctic krill form one of the largest biomasses of any individual animal species.BOOK, Nicol S, Endo Y,weblink Fisheries Technical Paper 367: Krill Fisheries of the World, Food and Agriculture Organization, FAO, 1997, File:Fungus, Minnowburn - - 1008404.jpg|It has been claimed that fungi make up 25% of the global biomass

Global rate of production

(File:Seawifs global biosphere.jpg|thumb|300px|right|Globally, terrestrial and oceanic habitats produce a similar amount of new biomass each year (56.4 billion tonnes C terrestrial and 48.5 billion tonnes C oceanic).)Net primary production is the rate at which new biomass is generated, mainly due to photosynthesis. Global primary production can be estimated from satellite observations. Satellites scan the normalised difference vegetation index (NDVI) over terrestrial habitats, and scan sea-surface chlorophyll levels over oceans. This results in 56.4 billion tonnes C/yr (53.8%), for terrestrial primary production, and 48.5 billion tonnes C/yr for oceanic primary production.JOURNAL, Field CB, Behrenfeld MJ, Randerson JT, Falkowski P, Primary production of the biosphere: integrating terrestrial and oceanic components, Science, 281, 5374, 237–40, July 1998, 9657713, 10.1126/science.281.5374.237, 1998Sci...281..237F,weblink Thus, the total photoautotrophic primary production for the Earth is about 104.9 billion tonnes C/yr. This translates to about 426 gC/m²/yr for land production (excluding areas with permanent ice cover), and 140 gC/m²/yr for the oceans.However, there is a much more significant difference in standing stocks—while accounting for almost half of total annual production, oceanic autotrophs account for only about 0.2% of the total biomass. Autotrophs may have the highest global proportion of biomass, but they are closely rivaled or surpassed by microbes.JOURNAL, Whitman WB, Coleman DC, Wiebe WJ, Prokaryotes: the unseen majority, Proceedings of the National Academy of Sciences of the United States of America, 95, 12, 6578–83, June 1998, 9618454, 33863, 10.1073/pnas.95.12.6578,weblink 1998PNAS...95.6578W, BOOK, Groombridge B, Jenkins M, World Atlas of Biodiversity: Earth's Living Resources in the 21st Century, BMC Public Health, 12, 439, World Conservation Monitoring Centre, United Nations Environment Programme, 2002,weblink 978-0-520-23668-4, 3408371, 22709383, 10.1186/1471-2458-12-439, Terrestrial freshwater ecosystems generate about 1.5% of the global net primary production.BOOK, Alexander, David E., vanc, Encyclopedia of Environmental Science, Springer Science+Business Media, Springer, 1 May 1999, 978-0-412-74050-3, Some global producers of biomass in order of productivity rates are{| class="wikitable sortable" border="1"! Producer! Biomass productivity(gC/m²/yr)! Ref! Total area(million km²)! Ref! Total production(billion tonnes C/yr)| Swamps and marshes
| Tropical rainforests
| Coral reefs
2,000| 0.28| Mark Spalding, Corinna Ravilious, and Edmund Green. 2001. World Atlas of Coral Reefs. Berkeley, California: University of California Press and UNEP/WCMC. 0.56
Algae>Algal beds 2,000| |||
Estuary>River estuaries 1,800| |||
| Temperate forests
1,250| 19| 24
Land use statistics by country>Cultivated lands 650 NAME-LIST-FORMAT = VANC YEAR= 2001 PUBLISHER= ROUTLEDGE URL= HTTPS://BOOKS.GOOGLE.COM/?ID=EW3MBJBW4OAC&PG=PA564, 978-0-415-21770-5, 17| 11
| Tundras
140| |||
Oceanic zone>Open ocean 125| 311| 39
| Deserts
3| 50| 0.15

See also

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Further reading

  • JOURNAL, Foley JA, Monfreda C, Ramankutty N, Zaks D, Our share of the planetary pie, Proceedings of the National Academy of Sciences of the United States of America, 104, 31, 12585–6, July 2007, 17646656, 1937509, 10.1073/pnas.0705190104, 2007PNAS..10412585F,
  • JOURNAL, Marina Fischer-Kowalski, Haberl H, Erb KH, Krausmann F, Gaube V, Bondeau A, Plutzar C, Gingrich S, Lucht W, Fischer-Kowalski M, Quantifying and mapping the human appropriation of net primary production in earth's terrestrial ecosystems, Proceedings of the National Academy of Sciences of the United States of America, 104, 31, 12942–7, July 2007, 17616580, 1911196, 10.1073/pnas.0704243104, 2007PNAS..10412942H,
  • BOOK, Purves, William K, Orians, Gordon H, vanc, 2007, Life: The Science of Biology, 8th, W. H. Freeman, 978-1-4292-0877-2,

External links

{{Wiktionary|biomass}} {{modelling ecosystems}}{{aquatic ecosystem topics|expanded=none}}

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