Biomass (ecology)

Short description: Total mass of living organisms in a given area (all species or selected species)

Apart from bacteria, the total global live biomass has been estimated as 550 or 560 thousand million tonnes C,[1] most of which is found in forests.[2]

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.[3]

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.[4] 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 (5.5-5.6×1011) tonnes C,[1][5] and the total annual primary production of biomass is just over 100 billion tonnes C/yr.[6] The total live biomass of bacteria may be as much as that of plants and animals[7] or may be much less.[1][8][9][10][11] The total number of DNA base pairs on Earth, as a possible approximation of global biodiversity, is estimated at (5.3±3.6)×1037, and weighs 50 billion tonnes.[12][13] Around 2020, anthropogenic mass (human-made material) is expected to exceed all living biomass on earth.[14]

## Contents

* 1 Ecological pyramids
* 2 Terrestrial biomass
* 3 Ocean biomass
* 4 Bacterial biomass
* 5 Global biomass
* 6 Global rate of production
* 7 See also
* 8 References
* 9 Further reading
* 10 External links

## Ecological pyramids

An energy pyramid illustrates how much energy is needed as it flows upward to support the next trophic level. Only about 10% of the energy transferred between each trophic level is converted to biomass.

Main page: Earth: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.

## Terrestrial biomass

Relative terrestrial biomasses
of vertebrates versus arthropods

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.

Changes in plant species in the terrestrial ecosystem can result in changes in the biomass of soil decomposer communities.[15] Biomass in C3 and C4 plant species can change in response to altered concentrations of CO2.[16] C3 plant species have been observed to increase in biomass in response to increasing concentrations of CO2 of up to 900 ppm.[17]

## Ocean biomass

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 fish

Ocean food web showing a network of food chains

Biomass pyramids
Compared to terrestrial biomass pyramids, aquatic pyramids are inverted at the base

Prochlorococcus, an influential bacterium

Phytoplankton 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 zooplankton that range in size from a few micrometers in diameter in the case of protistan microzooplankton to macroscopic gelatinous and crustacean 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.

A fourth trophic level can consist 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 a 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 which are r-strategists that grow and reproduce rapidly, so a small mass can have a fast rate of primary production. In contrast, terrestrial primary producers, such as forests, are K-strategists that grow and reproduce slowly, so a much larger mass is needed to achieve the same rate of primary production.

Among the phytoplankton at the base of the marine food web are members from a phylum of bacteria called cyanobacteria. Marine cyanobacteria include the smallest known photosynthetic organisms. The smallest of all, Prochlorococcus, is just 0.5 to 0.8 micrometres across.[18] In terms of individual numbers, Prochlorococcus is possibly the most plentiful species on Earth: a single millilitre of surface seawater can contain 100,000 cells or more. Worldwide, there are estimated to be several octillion (1027) individuals.[19] Prochlorococcus is ubiquitous between 40°N and 40°S and dominates in the oligotrophic (nutrient poor) regions of the oceans.[20] 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.[21]

## 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,[7] 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 2012[8] 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,[8] 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.[1] A census by the Deep Carbon Observatory project published in December 2018 gives a smaller figure of up to 23 billion tonnes of carbon.[9][10][11]

Geographic location  | Number of cells (× 1029)  | Billions of tonnes of carbon

|
|

Ocean floor

2.9[8] to 50[22]

4.1[8]

Open ocean

1.2[7]

1.7[7][8] to 10[7]

Terrestrial soil

2.6[7]

3.7[7][8] to 22[7]

Subsurface terrestrial

2.5 to 25[7]

3.5[7][8] to 215[7]

## Global biomass

External image Visualizing the biomass of life

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.[5][1] Most of this biomass is found on land, with only 5 to 10 billion tonnes C found in the oceans.[5] On land, there is about 1,000 times more plant biomass (phytomass) than animal biomass (zoomass).[23] About 18% of this plant biomass is eaten by the land animals.[24] However, in the ocean, the animal biomass is nearly 30 times larger than the plant biomass.[25][dubious - discuss] Most ocean plant biomass is eaten by the ocean animals.[24]

| 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

|
|
|
|
|
|
|
|
|

Terrestrial  |

Humans

2019

7.7 billion[26]

50 kg
(incl children)

30%

4.015×1036[27]

105

385

2005

4.63 billion adults

62 kg
(excl. children)[28]

|  |  |  |

287[28]

Cattle

|  |

1.3 billion[29]

400 kg

30%

|  |

156

520

Sheep and goats

2002

1.75 billion[30]

60 kg

30%

|  |

31.5

105

Chickens

|  |

24 billion

2 kg

30%

|  |

14.4

Ants

12,649[31]

|  |

107–108 billion[32]

3×10−6 kg
(0.003 grams)

30%

|  |

10–100

30-300

Earthworms

>7,000

1881
Darwin

1.3×106 billion[33]

3 g

30%[34]

|  |

1,140–2,280[33]

3,800–7,600[33]

Termites

>2,800

1996

|  |  |  |  |  |

445[35]

Marine

Blue whales[36]

Pre-whaling

340,000

|  |

40%[37]

|  |  |

2001

4,700

|  |

40%[37]

|  |  |

0.5

Fish

>10,000

2009

|  |  |  |  |  |

800-2,000[38]

Antarctic krill

1924–2004

7.8×1014

0.486 g

|  |  |  |

379[39]

Copepods
(a zooplankton)

13,000

|  |  |

10−6–10−9 kg

|  |

1×1037[40]

|  |

Cyanobacteria
(a picoplankton)

2003

|  |  |  |  |  |

1,000[41]

Global  |

Prokaryotes
(bacteria)

2018

1×1031 cells[1]

|  |  |  |

23,000[9] – 70,000[1]

Humans compose about 100 million tonnes of the Earth's dry biomass,[42] domesticated animals about 700 million tonnes, earthworms over 1,100 million tonnes,[33] and annual cereal crops about 2.3 billion tonnes.[43]

Biomass by life form

Humans and their livestock represent 96% of all mammals on earth in terms of biomass, whereas all wild mammals represent only 4%.[1]

The most successful animal species, in terms of biomass, may well be Antarctic krill, Euphausia superba, with a fresh biomass approaching 500 million tonnes.[39][44][45] As a group, the family of lanternfish are among the most populous vertebrates, with some estimates suggesting that they may have a total global biomass of 550–660 million tonnes, accounting for up to 65% of all deep-sea fish biomass.[46] However, as a group, the small aquatic invertebrates called copepods may form the largest animal biomass on earth.[47] 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.[48][49] It has been estimated that about 1% of the global biomass is due to phytoplankton.[50]

* Grasses, trees and shrubs have a much higher biomass than the animals that consume them

* Antarctic krill form one of the largest biomasses of any individual animal species.[44]

According to a 2020 study published in Nature, human-made materials, or anthropogenic mass, outweigh all living biomass on earth, with plastic alone exceeding the mass of all land and marine animals combined.[51][14]

## Global rate of production

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.[6] 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.[52][53]

Terrestrial freshwater ecosystems generate about 1.5% of the global net primary production.[54]

Some global producers of biomass in order of productivity rates are

Producer  | Biomass productivity
(gC/m2/yr)  | Ref  | Total area
(million km2)  | Ref  | Total production
(billion tonnes C/yr)

|
|
|
|
|

Swamps and marshes | 2,500  | [3] | 5.7  | [55] |
Tropical rainforests | 2,000  | [56] | 8  |  | 16
Coral reefs | 2,000  | [3] | 0.28  | [57] | 0.56
Algal beds | 2,000  | [3] |  |  |
River estuaries | 1,800  | [3] |  |  |
Temperate forests | 1,250  | [3] | 19  |  | 24
Cultivated lands  | 650  | [3][58] | 17  |  | 11
Tundras | 140  | [3][58] | 11.5-29.8  | [59][60] |
Open ocean | 125  | [3][58] | 311  |  | 39
Deserts | 3  | [58] | 50  |  | 0.15

## See also

* Biology:Biomass - Biological material from either living (see ecology) or recently living organisms (see bioenergy)
* Biology:Biomass (energy) - Biological material used as a renewable energy source
* Biomass partitioning
* Chemistry:Organic matter - Matter composed of organic compounds
* Earth:Productivity (ecology) - Rate of generation of biomass in an ecosystem
* Biology:Primary nutritional groups - Group of organisms
* Standing stock - Measurement of population per unit area or unit volume

## References

1. ↑ 1.0 1.1 1.2 1.3 1.4 1.5 1.6 1.7 "The biomass distribution on Earth". Proceedings of the National Academy of Sciences of the United States of America 115 (25): 6506–6511. June 2018. doi:10.1073/pnas.1711842115. PMID 29784790. Bibcode: 2018PNAS..115.6506B.
2. ↑ "Biomass". http://www.seps.sk/zp/fond/dieret/biomass.htm.
3. ↑ 3.0 3.1 3.2 3.3 3.4 3.5 3.6 3.7 3.8 Ricklefs, Robert E.; Miller, Gary Leon (2000). Ecology (4th ed.). Macmillan. p. 192. ISBN 978-0-7167-2829-0. https://books.google.com/books?id=6TMvdZQiySoC&q=temperate+forest+ecology+"net+primary+production"&pg=PA192.
4. ↑ IUPAC, Compendium of Chemical Terminology, 2nd ed. (the "Gold Book") (1997). Online corrected version:  (2006-) "biomass". doi:10.1351/goldbook.B00660
5. ↑ 5.0 5.1 5.2 Groombridge B, Jenkins MD (2000) Global biodiversity: Earth’s living resources in the 21st century Page 11. World Conservation Monitoring Centre, World Conservation Press, Cambridge
6. ↑ 6.0 6.1 "Primary production of the biosphere: integrating terrestrial and oceanic components". Science 281 (5374): 237–40. July 1998. doi:10.1126/science.281.5374.237. PMID 9657713. Bibcode: 1998Sci...281..237F. http://www.escholarship.org/uc/item/9gm7074q.
7. ↑ 7.00 7.01 7.02 7.03 7.04 7.05 7.06 7.07 7.08 7.09 7.10 "Prokaryotes: the unseen majority". Proceedings of the National Academy of Sciences of the United States of America 95 (12): 6578–83. June 1998. doi:10.1073/pnas.95.12.6578. PMID 9618454. PMC 33863. Bibcode: 1998PNAS...95.6578W. http://www.pnas.org/cgi/reprint/95/12/6578.pdf.
8. ↑ 8.0 8.1 8.2 8.3 8.4 8.5 8.6 8.7 "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. doi:10.1073/pnas.1203849109. PMID 22927371. Bibcode: 2012PNAS..10916213K.
9. ↑ 9.0 9.1 9.2 Deep Carbon Observatory (10 December 2018). "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". EurekAlert!. https://www.eurekalert.org/pub_releases/2018-12/tca-lid120318.php.
10. ↑ 10.0 10.1 Dockrill, Peter (11 December 2018). "Scientists Reveal a Massive Biosphere of Life Hidden Under Earth's Surface". Science Alert. https://www.sciencealert.com/scientists-lift-lid-on-massive-biosphere-of-life-hidden-under-earth-s-surface.
11. ↑ 11.0 11.1 Gabbatiss, Josh (11 December 2018). "Massive 'deep life' study reveals billions of tonnes of microbes living far beneath Earth's surface". The Independent. https://www.independent.co.uk/news/science/deep-life-microbes-underground-bacteria-earth-surface-carbon-observatory-science-study-a8677521.html.
12. ↑ "An Estimate of the Total DNA in the Biosphere". PLOS Biology 13 (6): e1002168. June 2015. doi:10.1371/journal.pbio.1002168. PMID 26066900.
13. ↑ Nuwer, Rachel (18 July 2015). "Counting All the DNA on Earth". The New York Times (New York). ISSN 0362-4331. https://www.nytimes.com/2015/07/21/science/counting-all-the-dna-on-earth.html.
14. ↑ 14.0 14.1 Elhacham, Emily et al. (2020). "Global human-made mass exceeds all living biomass". Nature 588 (7838): 442–444. doi:10.1038/s41586-020-3010-5. PMID 33299177. Bibcode: 2020Natur.588..442E.
15. ↑ Spehn, Eva M.; Joshi, Jasmin; Schmid, Bernhard; Alphei, Jörn; Körner, Christian (2000). "Plant diversity effects on soil heterotrophic activity in experimental grassland ecosystems". Plant and Soil 224 (2): 217–230. doi:10.1023/A:1004891807664. http://link.springer.com/10.1023/A:1004891807664.
16. ↑ He, Jin-Sheng; Bazzaz, Fakhri A.; Schmid, Bernhard (2002). "Interactive Effects of Diversity, Nutrients and Elevated CO2 on Experimental Plant Communities". Oikos 97 (3): 337–348. doi:10.1034/j.1600-0706.2002.970304.x. ISSN 0030-1299. https://www.jstor.org/stable/3547655.
17. ↑ Drag, David W; Slattery, Rebecca; Siebers, Matthew; DeLucia, Evan H; Ort, Donald R; Bernacchi, Carl J (2020-03-12). "Soybean photosynthetic and biomass responses to carbon dioxide concentrations ranging from pre-industrial to the distant future". Journal of Experimental Botany (Oxford University Press (OUP)) 71 (12): 3690–3700. doi:10.1093/jxb/eraa133. ISSN 0022-0957. PMID 32170296.
18. ↑ "Patterns and implications of gene gain and loss in the evolution of Prochlorococcus". PLOS Genetics 3 (12): e231. December 2007. doi:10.1371/journal.pgen.0030231. PMID 18159947.
19. ↑ Nemiroff, R.; Bonnell, J., eds (27 September 2006). "Earth from Saturn". Astronomy Picture of the Day. NASA. https://apod.nasa.gov/apod/ap060927.html.
20. ↑ "Prochlorococcus, a marine photosynthetic prokaryote of global significance". Microbiology and Molecular Biology Reviews 63 (1): 106–27. March 1999. doi:10.1128/MMBR.63.1.106-127.1999. PMID 10066832.
21. ↑ "The Most Important Microbe You've Never Heard Of". https://www.npr.org/templates/story/story.php?storyId=91448837.
22. ↑ "Significant contribution of Archaea to extant biomass in marine subsurface sediments". Nature 454 (7207): 991–994. August 2008. doi:10.1038/nature07174. PMID 18641632. Bibcode: 2008Natur.454..991L.
23. ↑ Gosh, Iman (20 August 2021). "Misc All the Biomass of Earth, in One Graphic". Visual Capitalist. https://www.visualcapitalist.com/all-the-biomass-of-earth-in-one-graphic/.
24. ↑ 24.0 24.1 Hartley, Sue (2010) The 300 Million Years War: Plant Biomass v Herbivores Royal Institution Christmas Lecture.
25. ↑ Darlington, P (1966) http://encyclopedia2.thefreedictionary.com/Terrestrial+Fauna "Biogeografia". Published in The Great Soviet Encyclopedia, 3rd Edition (1970–1979).
26. ↑ "world population clock". https://www.worldometers.info/world-population/.
27. ↑ Freitas, Robert A. Jr.Nanomedicine 3.1 Human Body Chemical Composition Foresight Institute, 1998
28. ↑ 28.0 28.1 "The weight of nations: an estimation of adult human biomass". BMC Public Health 12 (1): 439. June 2012. doi:10.1186/1471-2458-12-439. PMID 22709383.
29. ↑ Cattle Today. "Breeds of Cattle at CATTLE TODAY". Cattle-today.com. http://cattle-today.com/.
30. ↑ World's Rangelands Deteriorating Under Mounting Pressure Earth Policy Institute 2002
31. ↑ "Archived copy". http://osuc.biosci.ohio-state.edu/hymenoptera/tsa.sppcount?the_taxon=Formicidae.
32. ↑ Embery, Joan; Lucaire, Ed; Karel, Havlicek (1983). Joan Embery's collection of amazing animal facts. New York: Delacorte Press. ISBN 978-0-385-28486-8. https://archive.org/details/joanemberyscolle0000embe.
33. ↑ 33.0 33.1 33.2 33.3 "Darwin's win-win for Global Worming?". 2017. https://vermecology.wordpress.com/2017/02/12/nature-article-to-commemorate-charles-darwins-birthday-on-12th-feb/.
34. ↑ Earthworms: their ecology and relationships with soils and land use. Sydney: Academic Press. 1985. ISBN 978-0-12-440860-9.
35. ↑ Sum of [(biomass m−22)*(area m2)] 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
36. ↑ Humphries, Stuart, ed (August 2010). "The impact of whaling on the ocean carbon cycle: why bigger was better". PLOS ONE 5 (8): e12444. doi:10.1371/journal.pone.0012444. PMID 20865156. Bibcode: 2010PLoSO...512444P. (Table 1)
37. ↑ 37.0 37.1 "Whaling and Deep-Sea Biodiversity". Conservation Biology 10 (2): 653–654. 1996. doi:10.1046/j.1523-1739.1996.10020653.x.
38. ↑ Wilson, R. W.; Millero, F. J.; Taylor, J. R.; Walsh, P. J.; Christensen, V.; Jennings, S.; Grosell, M. (16 January 2009). "Contribution of Fish to the Marine Inorganic Carbon Cycle". Science 323 (5912): 359–362. doi:10.1126/science.1157972. PMID 19150840. Bibcode: 2009Sci...323..359W. (This article provides a first estimate of global fish "wet weight" biomass)
39. ↑ 39.0 39.1 "A re-appraisal of the total biomass and annual production of Antarctic krill". Deep-Sea Research Part I 56 (5): 727–740. 2009. doi:10.1016/j.dsr.2008.12.007. Bibcode: 2009DSRI...56..727A. http://www.iced.ac.uk/documents/Atkinson et al, Deep Sea Research I, 2009.pdf.
40. ↑ "Biogeochemical fluxes through mesozooplankton". Global Biogeochemical Cycles 20 (2): 2003. 2006. doi:10.1029/2005GB002511. Bibcode: 2006GBioC..20.2003B. http://europa.agu.org/?view=article&uri=/journals/gb/gb0602/2005GB002511/2005GB002511.xml.
41. ↑ "Estimates of global cyanobacterial biomass and its distribution". Algological Studies 109: 213–217. 2003. doi:10.1127/1864-1318/2003/0109-0213. http://sbsc.wr.usgs.gov/products/pdfs/GarciaPichel_et_al_2003_Estimates_of_global_cyanobacterial.pdf.
42. ↑ 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.[clarification needed]
43. ↑ FAO Statistical Yearbook 2013: page 130 - http://www.fao.org/docrep/018/i3107e/i3107e.PDF
44. ↑ 44.0 44.1 Fisheries Technical Paper 367: Krill Fisheries of the World. FAO. 1997. http://www.fao.org/documents/show_cdr.asp?url_file=//DOCREP/003/W5911E/w5911e00.htm.
45. ↑ Ross, R. M. and Quetin, L. B. (1988). Euphausia superba: a critical review of annual production. Comp. Biochem. Physiol. 90B, 499-505.
46. ↑ Schwarzhans, Werner; Carnevale, Giorgio (2021-03-19). "The rise to dominance of lanternfishes (Teleostei: Myctophidae) in the oceanic ecosystems: a paleontological perspective" (in en). Paleobiology 47 (3): 446–463. doi:10.1017/pab.2021.2. ISSN 0094-8373.
47. ↑ "Biology of Copepods". Carl von Ossietzky University of Oldenburg. http://www.uni-oldenburg.de/zoomorphology/Biology.html.
48. ↑ "Contribution of fish to the marine inorganic carbon cycle". Science 323 (5912): 359–362. January 2009. doi:10.1126/science.1157972. PMID 19150840. Bibcode: 2009Sci...323..359W.
49. ↑ Researcher gives first-ever estimate of worldwide fish biomass and impact on climate change PhysOrg.com, 15 January 2009.
50. ↑ "Cell death in planktonic, photosynthetic microorganisms". Nature Reviews. Microbiology 2 (8): 643–655. August 2004. doi:10.1038/nrmicro956. PMID 15263899.
51. ↑ Laville, Sandra (December 9, 2020). "Human-made materials now outweigh Earth's entire biomass – study". The Guardian. https://www.theguardian.com/environment/2020/dec/09/human-made-materials-now-outweigh-earths-entire-biomass-study.
52. ↑ "Prokaryotes: the unseen majority". Proceedings of the National Academy of Sciences of the United States of America 95 (12): 6578–83. June 1998. doi:10.1073/pnas.95.12.6578. PMID 9618454. Bibcode: 1998PNAS...95.6578W.
53. ↑ World Atlas of Biodiversity: Earth's Living Resources in the 21st Century. 12. World Conservation Monitoring Centre, United Nations Environment Programme. 2002. 439. doi:10.1186/1471-2458-12-439. ISBN 978-0-520-23668-4. https://books.google.com/books?id=_kHeAXV5-XwC&q=biomass.
54. ↑ Alexander, David E. (1 May 1999). Encyclopedia of Environmental Science. Springer. ISBN 978-0-412-74050-3.
55. ↑ https://www.ramsar.org/sites/default/files/documents/library/info2007-01-e.pdf [bare URL PDF]
56. ↑ Ricklefs, Robert E.; Miller, Gary Leon (2000). Ecology (4th ed.). Macmillan. p. 197. ISBN 978-0-7167-2829-0. https://books.google.com/books?id=6TMvdZQiySoC&q=primary+production+biomass+g+m+yr&pg=PA197.
57. ↑ Mark Spalding, Corinna Ravilious, and Edmund Green. 2001. World Atlas of Coral Reefs. Berkeley, California: University of California Press and UNEP/WCMC.
58. ↑ 58.0 58.1 58.2 58.3 Park, Chris C. (2001). The environment: principles and applications (2nd ed.). Routledge. p. 564. ISBN 978-0-415-21770-5. https://books.google.com/books?id=Ew3MBjbw4OAC&pg=PA564.
59. ↑ "Tundra - Biomes - WWF". https://www.worldwildlife.org/biomes/tundra.
60. ↑ "Tundra". 2020-01-17. https://storymaps.arcgis.com/stories/93e3669fa9ab42c0b98e3c8ad31f25f6. "the tundra is a vast and treeless land which covers about 20% of the Earth's surface, circumnavigating the North pole."

## Further reading

* "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. doi:10.1073/pnas.0705190104. PMID 17646656. Bibcode: 2007PNAS..10412585F.
* "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. doi:10.1073/pnas.0704243104. PMID 17616580. Bibcode: 2007PNAS..10412942H.
* Purves, William K; Orians, Gordon H (2007). Life: The Science of Biology (8th ed.). W. H. Freeman. ISBN 978-1-4292-0877-2.

## External links

* The mass of all life on Earth is staggering — until you consider how much we’ve lost
* Counting bacteria
* Trophic levels
* Biomass distributions for high trophic-level fishes in the North Atlantic, 1900–2000

* v
* t
* e

Ecology: Modelling ecosystems: Trophic components

General|

* Abiotic component
* Abiotic stress
* Behaviour
* Biogeochemical cycle
* Biomass
* Biotic component
* Biotic stress
* Carrying capacity
* Competition
* Ecosystem
* Ecosystem ecology
* Ecosystem model
* Keystone species
* List of feeding behaviours
* Metabolic theory of ecology
* Productivity
* Resource

Producers|

* Autotrophs
* Chemosynthesis
* Chemotrophs
* Foundation species
* Mixotrophs
* Myco-heterotrophy
* Mycotroph
* Organotrophs
* Photoheterotrophs
* Photosynthesis
* Photosynthetic efficiency
* Phototrophs
* Primary nutritional groups
* Primary production

Consumers|

* Apex predator
* Bacterivore
* Carnivores
* Chemoorganotroph
* Foraging
* Generalist and specialist species
* Intraguild predation
* Herbivores
* Heterotroph
* Heterotrophic nutrition
* Insectivore
* Mesopredators
* Mesopredator release hypothesis
* Omnivores
* Optimal foraging theory
* Planktivore
* Predation
* Prey switching

Decomposers|

* Chemoorganoheterotrophy
* Decomposition
* Detritivores
* Detritus

Microorganisms|

* Archaea
* Bacteriophage
* Environmental microbiology
* Lithoautotroph
* Lithotrophy
* Microbial cooperation
* Microbial ecology
* Microbial food web
* Microbial intelligence
* Microbial loop
* Microbial mat
* Microbial metabolism
* Phage ecology

Food webs|

* Biomagnification
* Ecological efficiency
* Ecological pyramid
* Energy flow
* Food chain
* Trophic level

Example webs|

* Cold seeps
* Hydrothermal vents
* Intertidal
* Kelp forests
* Lakes
* North Pacific Subtropical Gyre
* Rivers
* San Francisco Estuary
* Soil
* Tide pool

Processes|

* Ascendency
* Bioaccumulation
* Cascade effect
* Climax community
* Competitive exclusion principle
* Consumer-resource systems
* Copiotrophs
* Dominance
* Ecological network
* Ecological succession
* Energy quality
* Energy Systems Language
* f-ratio
* Feed conversion ratio
* Feeding frenzy
* Mesotrophic soil
* Nutrient cycle
* Oligotroph
* Paradox of the plankton
* Trophic cascade
* Trophic mutualism
* Trophic state index

Defense,
counter|

* Animal coloration
* Antipredator adaptations
* Camouflage
* Deimatic behaviour
* Herbivore adaptations to plant defense
* Mimicry
* Plant defense against herbivory
* Predator avoidance in schooling fish

* v
* t
* e

Ecology: Modelling ecosystems: Other components

Population
ecology|

* Abundance
* Allee effect
* Depensation
* Ecological yield
* Effective population size
* Intraspecific competition
* Logistic function
* Malthusian growth model
* Maximum sustainable yield
* Overpopulation in wild animals
* Overexploitation
* Population cycle
* Population dynamics
* Population modeling
* Population size
* Predator–prey (Lotka–Volterra) equations
* Recruitment
* Resilience
* Small population size
* Stability

Species|

* Biodiversity
* Density-dependent inhibition
* Ecological effects of biodiversity
* Ecological extinction
* Endemic species
* Flagship species
* Gradient analysis
* Indicator species
* Introduced species
* Invasive species
* Latitudinal gradients in species diversity
* Minimum viable population
* Neutral theory
* Occupancy–abundance relationship
* Population viability analysis
* Priority effect
* Rapoport's rule
* Relative abundance distribution
* Relative species abundance
* Species diversity
* Species homogeneity
* Species richness
* Species distribution
* Species-area curve
* Umbrella species

Species
interaction|

* Antibiosis
* Biological interaction
* Commensalism
* Community ecology
* Ecological facilitation
* Interspecific competition
* Mutualism
* Parasitism
* Storage effect
* Symbiosis

Spatial
ecology|

* Biogeography
* Cross-boundary subsidy
* Ecocline
* Ecotone
* Ecotype
* Disturbance
* Edge effects
* Foster's rule
* Habitat fragmentation
* Ideal free distribution
* Intermediate Disturbance Hypothesis
* Island biogeography
* Landscape ecology
* Landscape epidemiology
* Landscape limnology
* Metapopulation
* Patch dynamics
* r/K selection theory
* Resource selection function
* Source–sink dynamics

Niche|

* Ecological niche
* Ecological trap
* Ecosystem engineer
* Environmental niche modelling
* Guild
* Habitat
* Marine habitats
* Limiting similarity
* Niche apportionment models
* Niche construction
* Niche differentiation

Other
networks|

* Assembly rules
* Bateman's principle
* Bioluminescence
* Ecological collapse
* Ecological debt
* Ecological deficit
* Ecological energetics
* Ecological indicator
* Ecological threshold
* Ecosystem diversity
* Emergence
* Extinction debt
* Kleiber's law
* Liebig's law of the minimum
* Marginal value theorem
* Thorson's rule
* Xerosere

Other|

* Allometry
* Alternative stable state
* Balance of nature
* Biological data visualization
* Ecocline
* Ecological economics
* Ecological footprint
* Ecological forecasting
* Ecological humanities
* Ecological stoichiometry
* Ecopath
* Ecosystem based fisheries
* Endolith
* Evolutionary ecology
* Functional ecology
* Industrial ecology
* Macroecology
* Microecosystem
* Natural environment
* Regime shift
* Systems ecology
* Urban ecology
* Theoretical ecology

List of ecology topics

* v
* t
* e

Aquatic ecosystems

Aquatic ecosystems – general and freshwater components

| General|

* Acoustic ecology
* Adaptation
* Agent-based models
* Algal bloom
* Anoxic waters
* Aquatic animals (Insects
* Mammals)
* Aquatic plants
* Aquatic science
* Benthos
* Biodiversity research
* Bioluminescence
* Biomass
* Biomonitoring
* Cascade effect
* Colored dissolved organic matter
* Camouflage and mimicry
* Dead zone
* Ecohydrology
* Ecosystems
* Eutrophication
* Fisheries science
* Food chain
* Food web
* GIS and aquatic science
* Hydrobiology
* Hypoxia
* Isotope analysis
* Microbial ecology
* Microbial food web
* Microbial loop
* Nekton
* Neuston
* Particle
* Pelagic zone
* Photic zone
* Phytoplankton
* Plankton
* Pleuston
* Predation
* Productivity
* Ramsar Convention
* Respiration
* Schooling
* Sediment trap
* Siltation
* Spawning
* Substrate
* Thermal pollution
* Toxicology
* Trophic level
* Water column
* Zooplankton
* More...

|
|

Freshwater|

* Biology
* Biomes
* Ecosystems
* freshwater
* lake
* river
* Fish
* Hyporheic zone
* Limnology
* Lake stratification
* Macrophyte
* Pond
* Fish pond
* Rheotaxis
* Stream bed
* Stream pool
* Trophic state index
* Upland and lowland
* Water garden
* Wetland
* brackish marsh
* freshwater marsh
* swamp
* bog
* fen
* Environmental quality
* More...

Ecoregions|

* Freshwater (List)
* Marine (List)
* The Everglades
* Maharashtra
* The North Pacific Subtropical Gyre
* The San Francisco Estuary

Aquatic ecosystems – marine components

| Marine|

* Marine biology
* Marine chemistry
* Deep scattering layer
* Diel vertical migration
* Ecosystems
* large marine
* marine)
* f-ratio
* Iron fertilization
* Marine snow
* Ocean nourishment
* Oceanic physical-biological process
* Ocean turbidity
* Photophore
* Thorson's rule
* Upwelling
* Whale fall
* More...

|
|

Marine
life|

* Bacteriophages
* Census
* Fish
* coastal
* coral reef
* deep sea
* demersal
* pelagic
* Deep sea communities
* Deep sea creature
* Deep-water coral
* Invertebrates
* Larvae
* Mammals
* Marine life
* Algae and plants
* Microorganisms
* Paradox of the plankton
* Reptiles
* Seabirds
* Seashore wildlife
* Vertebrates
* Wild fisheries

Marine
habitats|

* Bay mud
* Coastal biogeomorphology
* Cold seeps
* Coral reefs
* Davidson Seamount
* Estuaries
* Intertidal ecology
* Intertidal wetlands
* Kelp forests
* Hydrothermal vents
* Lagoons
* Mangroves
* Marine biomes
* Marine habitats
* Mudflats
* Rocky shores
* Salt marshes
* Salt pannes and pools
* Seagrass meadows
* Sponge grounds
* Sponge reefs
* Tide pools

Issues|

* Coral bleaching
* Ecological values of mangroves
* Fisheries and climate change
* HERMIONE
* Marine conservation
* Marine conservation activism
* Marine pollution
* Marine Protected Area

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