Biomass (ecology)

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

The total global live biomass has been estimated at about 550 billion tonnes carbon,[1] most of which is found in forests.

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.[2]

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.[3] 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.

In 2018, Bar-On et al. estimated the total live biomass on Earth at about 550 billion (5.5×1011) tonnes C,[1] most of it in plants. In 1998 Field et.al. estimated the total annual net primary production of biomass at just over 100 billion tonnes C/yr.[4] The total live biomass of bacteria was once thought to be about the same as plants,[5] but recent studies suggest it is significantly less.[1][6][7][8][9] 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.[10][11] Anthropogenic mass (human-made material) is expected to exceed all living biomass on earth at around the year 2020.[12]

## 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 article: 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.[13] Biomass in C3 and C4 plant species can change in response to altered concentrations of CO2.[14] C3 plant species have been observed to increase in biomass in response to increasing concentrations of CO2 of up to 900 ppm.[15]

## 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.[16] 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.[17] Prochlorococcus is ubiquitous between 40°N and 40°S and dominates in the oligotrophic (nutrient poor) regions of the oceans.[18] 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.[19]

## Bacterial biomass

Bacteria and archaea are both classified as prokaryotes, and their biomass is commonly estimated together. The global biomass of prokaryotes is estimated at about 30 billion tonnes C,[20] dominated by bacteria.[1]

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

|
|

Open ocean

1.2[1][5]

1.6[1] to 2.2[5]

Ocean subsurface

5[20]

10[20]

Terrestrial soil

3[1]

8[1]

Terrestrial subsurface

2 to 6[20]

4 to 12[20]

Total

11 to 15[20]

23 to 31[20]

The estimates for the global biomass of prokaryotes had changed significantly over recent decades, as more data became available. A much-cited study from 1998[5] collected data on abundances (number of cells) of bacteria and archaea in different natural environments, and estimated their total biomass at 350 to 550 billion tonnes C. This vast amount is similar to the biomass of carbon in all plants.[1][5] The vast majority of bacteria and archaea were estimated to be in sediments deep below the seafloor or in the deep terrestrial biosphere (in deep continental aquifers). However, updated measurements reported in a 2012 study[6] reduced the calculated prokaryotic biomass in deep subseafloor sediments from the original ≈300 billion tonnes C to ≈4 billion tonnes C (range 1.5-22 billion tonnes). This update originates from much lower estimates of both the prokaryotic abundance and their average weight.

A census published in PNAS in May 2018 estimated global bacterial biomass at ≈70 billion tonnes C, of which ≈60 billion tonnes are in the terrestrial deep subsurface.[1] It also estimated the global biomass of archaea at ≈7 billion tonnes C. A later study by the Deep Carbon Observatory published in 2018 reported a much larger dataset of measurements, and updated the total biomass estimate in the deep terrestrial biosphere. It used this new knowledge and previous estimates to update the global biomass of bacteria and archaea to 23 - 31 billion tonnes C.[20] Roughly 70% of the global biomass was estimated to be found in the deep subsurface.[7][21] The estimated number of prokaryotic cells globally was estimated to be 11-15 × 1029.[20] With this information, the authors of the May 2018 PNAS article[1] revised their estimate for the global biomass of prokaryotes to ≈30 billion tonnes C,[22] similar to the Deep Carbon Observatory estimate.[20]

These estimates convert global abundance of prokaryotes into global biomass using average cellular biomass figures that are based on limited data. Recent estimates used an average cellular biomass of about 20-30 femtogram carbon (fgC) per cell in the subsurface and terrestrial habitats.[1][20][23]

## Global biomass

External image Visualizing the biomass of life

The total global biomass has been estimated at about 550 billion tonnes C. [24][1] A breakdown of the global biomass is given by kingdom in the table below, based on a 2018 study by Bar-On et. al.[1]

Kingdom  | Global biomass in billion tonnes of carbon  | Global dry biomass in billion tonnes  | Global wet biomass in billion tonnes  | Image

|
|
|
|

Plantae

450[1]

900

2700

Bacteria \+ Archaea

30[20][22]

200

Fungi

12[1]

Protista

4[1]

Animalia

2[1]

Total

500

1000

3000

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

Animals represent less than 0.5% of the total biomass on Earth, with about 2 billion tonnes C in total. Most animal biomass is found in the oceans, where arthropods, such as copepods, account for about 1 billion tonnes C and fish for another 0.7 billion tonnes C.[1] Roughly half of the biomass of fish in the world are mesopelagic, such as lanternfish,[25] spending most of the day in the deep, dark waters.[26] Marine mammals such as whales and dolphins account for about 0.006 billion tonnes C.[27]

Land animals account for about 500 million tonnes C, or about 20% of the biomass of animals on Earth.[1] Terrestrial arthropods account for about 150 million tonnes C, most of which is found in the topsoil.[28] Land mammals account for about 180 million tonnes C, most of which are humans (about 80 million tonnes C) and domesticated mammals (about 90 million tonnes C). Wild terrestrial mammals account for only about 3 million tonnes C, less than 2% of the total mammalian biomass on land.[27]

The global biomass broken down by kingdom and into taxonomic groups for animals.[1] The estimates for bacteria and archaea have been updated to 30 billion tonnes C combined since this figure was made.[20]

Most of the global biomass is found on land, with only 5 to 10 billion tonnes C found in the oceans.[24] On land, there is about 1,000 times more plant biomass (phytomass) than animal biomass (zoomass).[29] About 18% of this plant biomass is eaten by the land animals.[30] However, marine animals eat most of the marine autotrophs, and the biomass of marine animals is greater than that of marine autotrophs. [1][30]

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.[31][12][32]

| name  | number of species  | date of estimate  | individual count  | mean living mass of individual  | percent biomass (dried)  | global dry biomass in million tonnes  | global wet (fresh) biomass in million tonnes

|
|
|
|
|
|
|
|

Terrestrial  |

Humans

November 2022

8 billion[33]

50 kg
(incl children)[34]

40%[35]

160

400[27]

2005

4.63 billion adults

62 kg
(excl. children)[36]

|  |  |

287[36]

Cattle

2021

1.5 billion[37]

300 kg

30%

125

416[27]

Sheep

2021

1.3 billion[37]

30 kg

30%

39[27]

Goats

2021

1.1 billion[37]

30 kg

30%

32[27]

Chickens

2021

26 billion

0.9 kg for broilers, 1.8 kg for layers[38]

30%

8[1]

Ants

15,700[39]

2022

20-90×1015[39]

3.7 mg[28]-5.5 mg[39]

22.8%[40]

10–100[39]

40-450

Earthworms

7,000-30,000[41]

2016

|  |

10 mg (dry weight)[42]

10-25%[43]

400[1]

1,600

Termites

2,972[citation needed]

2022

|  |

2 mg[40]

27%[40]

100[28][44]

440[45]

Nematodes

2019

4.4×1020[46]

|  |

20%[40]

| 60  |

300[46]

Marine

Blue whales[47]

Pre-whaling

340,000

|  |

40%[48]

|  |

2023

50,000[27]

60,000 kg

40%[48]

1.2

3[27]

Fish

>20,000[49]

2022

|  |  |

30%[50]

3,000

9,000[26]

Antarctic krill

2008

7.8×1014[51]

0.486 g[51]

|  |  |

379 (in peak season)[51]

Copepods
(a zooplankton)

13,000

|  |  |

10−6–10−9 kg

|  |  |

Cyanobacteria
(a picoplankton)

2003

|  |  |  |  |

1,000[52]

## 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.[4] 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.

Terrestrial freshwater ecosystems generate about 1.5% of the global net primary production.[53]

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  | [2] | 5.7  | [54] |
Tropical rainforests | 2,000  | [55] | 8  |  | 16
Coral reefs | 2,000  | [2] | 0.28  | [56] | 0.56
Algal beds | 2,000  | [2] |  |  |
River estuaries | 1,800  | [2] |  |  |
Temperate forests | 1,250  | [2] | 19  |  | 24
Cultivated lands  | 650  | [2][57] | 17  |  | 11
Tundras | 140  | [2][57] | 11.5-29.8  | [58][59] |
Open ocean | 125  | [2][57] | 311  |  | 39
Deserts | 3  | [57] | 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 size per unit area or unit volume
* Slash-and-burn
* Stubble burning

## References

1. ↑ 1.00 1.01 1.02 1.03 1.04 1.05 1.06 1.07 1.08 1.09 1.10 1.11 1.12 1.13 1.14 1.15 1.16 1.17 1.18 1.19 1.20 1.21 1.22 1.23 1.24 "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. ↑ 2.0 2.1 2.2 2.3 2.4 2.5 2.6 2.7 2.8 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.
3. ↑ IUPAC, Compendium of Chemical Terminology, 2nd ed. (the "Gold Book") (1997). Online corrected version:  (2006-) "biomass". doi:10.1351/goldbook.B00660
4. ↑ 4.0 4.1 "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.
5. ↑ 5.0 5.1 5.2 5.3 5.4 "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.
6. ↑ 6.0 6.1 "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.
7. ↑ 7.0 7.1 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.
8. ↑ 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.
9. ↑ 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.
10. ↑ "An Estimate of the Total DNA in the Biosphere". PLOS Biology 13 (6): e1002168. June 2015. doi:10.1371/journal.pbio.1002168. PMID 26066900.
11. ↑ 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.
12. ↑ 12.0 12.1 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.
13. ↑ 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.
14. ↑ 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.
15. ↑ 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.
16. ↑ "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.
17. ↑ Nemiroff, R.; Bonnell, J., eds (27 September 2006). "Earth from Saturn". Astronomy Picture of the Day. NASA. https://apod.nasa.gov/apod/ap060927.html.
18. ↑ "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.
19. ↑ "The Most Important Microbe You've Never Heard Of". https://www.npr.org/templates/story/story.php?storyId=91448837.
20. ↑ 20.00 20.01 20.02 20.03 20.04 20.05 20.06 20.07 20.08 20.09 20.10 20.11 20.12 Magnabosco, C.; Lin, L.-H.; Dong, H.; Bomberg, M.; Ghiorse, W.; Stan-Lotter, H.; Pedersen, K.; Kieft, T. L. et al. (24 September 2018). "The biomass and biodiversity of the continental subsurface" (in en). Nature Geoscience 11 (10): 707–717. doi:10.1038/s41561-018-0221-6. ISSN 1752-0908. Bibcode: 2018NatGe..11..707M. https://www.nature.com/articles/s41561-018-0221-6.
21. ↑ Observatory, Deep Carbon. "Life in deep Earth totals 15 to 23 billion tons of carbon—hundreds of times more than humans" (in en). https://phys.org/news/2018-12-life-deep-earth-totals-billion.html.
22. ↑ 22.0 22.1 Bar-On, Yinon M.; Milo, Ron (21 February 2019). "Towards a quantitative view of the global ubiquity of biofilms" (in en). Nature Reviews Microbiology 17 (4): 199–200. doi:10.1038/s41579-019-0162-0. ISSN 1740-1534. PMID 30792541. https://www.nature.com/articles/s41579-019-0162-0.
23. ↑ Griebler, Christian; Mindl, Birgit; Slezak, Doris; Geiger-Kaiser, Margot (2002-06-26). "Distribution patterns of attached and suspended bacteria in pristine and contaminated shallow aquifers studied with an in situ sediment exposure microcosm" (in en). Aquatic Microbial Ecology 28 (2): 117–129. doi:10.3354/ame028117. ISSN 0948-3055. https://www.int-res.com/abstracts/ame/v28/n2/p117-129/.
24. ↑ 24.0 24.1 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
25. ↑ 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. Bibcode: 2021Pbio...47..446S.
26. ↑ 26.0 26.1 Hatton, Ian A.; Heneghan, Ryan F.; Bar-On, Yinon M.; Galbraith, Eric D. (2021-11-12). "The global ocean size spectrum from bacteria to whales" (in en). Science Advances 7 (46): eabh3732. doi:10.1126/sciadv.abh3732. ISSN 2375-2548. PMID 34757796. Bibcode: 2021SciA....7.3732H.
27. ↑ 27.0 27.1 27.2 27.3 27.4 27.5 27.6 27.7 Greenspoon, Lior; Krieger, Eyal; Sender, Ron; Rosenberg, Yuval; Bar-On, Yinon M.; Moran, Uri; Antman, Tomer; Meiri, Shai et al. (2023-03-07). "The global biomass of wild mammals" (in en). Proceedings of the National Academy of Sciences 120 (10): e2204892120. doi:10.1073/pnas.2204892120. ISSN 0027-8424. PMID 36848563. Bibcode: 2023PNAS..12004892G.
28. ↑ 28.0 28.1 28.2 Rosenberg, Yuval; Bar-On, Yinon M.; Fromm, Amir; Ostikar, Meital; Shoshany, Aviv; Giz, Omer; Milo, Ron (3 Feb 2023). "The global biomass and number of terrestrial arthropods" (in en). Science Advances 9 (5): eabq4049. doi:10.1126/sciadv.abq4049. ISSN 2375-2548. PMID 36735788. Bibcode: 2023SciA....9.4049R.
29. ↑ 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/.
30. ↑ 30.0 30.1 Hartley, Sue (2010) The 300 Million Years War: Plant Biomass v Herbivores Royal Institution Christmas Lecture.
31. ↑ 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.
32. ↑ "Anthropogenic mass: Comparing human-made mass to the living Biomass on earth". http://anthropomass.org/.
33. ↑ Nations, United. "Day of 8 Billion" (in en). https://www.un.org/en/dayof8billion.
34. ↑ Hern, Warren M. (September 1999). "How Many Times Has the Human Population Doubled? Comparisons with Cancer". Population and Environment 21 (1): 59–80. doi:10.1007/BF02436121. https://www.jstor.org/stable/27503685.
35. ↑ Jéquier, E.; Constant, F. (February 2010). "Water as an essential nutrient: the physiological basis of hydration" (in en). European Journal of Clinical Nutrition 64 (2): 115–123. doi:10.1038/ejcn.2009.111. ISSN 1476-5640. PMID 19724292. https://www.nature.com/articles/ejcn2009111.
36. ↑ 36.0 36.1 "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.
37. ↑ 37.0 37.1 37.2 "FAOSTAT". https://www.fao.org/faostat/en/#data/QCL.
38. ↑ IPCC 2006, 2006 IPCC Guidelines for National Greenhouse Gas Inventories, Prepared by the National Greenhouse Gas Inventories Programme, Eggleston H.S., Buendia L., Miwa K., Ngara T. and Tanabe K. (eds). Published: IGES, Japan.
39. ↑ 39.0 39.1 39.2 39.3 Schultheiss, Patrick; Nooten, Sabine; Wang, Runxi; Wong, Mark; Brassard, François; Benoit, Guénard (September 19, 2022). "The abundance, biomass, and distribution of ants on Earth" (in en). Proceedings of the National Academy of Sciences 119 (40): e2201550119. doi:10.1073/pnas.2201550119. ISSN 0027-8424. PMID 36122199. Bibcode: 2022PNAS..11901550S.
40. ↑ 40.0 40.1 40.2 40.3 Petersen, Henning; Luxton, Malcolm (December 1982). "A Comparative Analysis of Soil Fauna Populations and Their Role in Decomposition Processes". Oikos 39 (3): 288–388. doi:10.2307/3544689. https://www.researchgate.net/publication/274752964. Retrieved July 26, 2023.
41. ↑ Joint Research Centre (European Commission); Johnson, Nancy C.; Scheu, Stefan; Ramirez, Kelly S.; Lemanceau, Philippe; Eggleton, Paul; Jones, Arwyn; Moreira, Fatima M. S. et al. (2016). Global soil biodiversity atlas. LU: Publications Office of the European Union. doi:10.2788/2613. ISBN 978-92-79-48168-0. https://data.europa.eu/doi/10.2788/2613.
42. ↑ Fierer, Noah; Strickland, Michael S.; Liptzin, Daniel; Bradford, Mark A.; Cleveland, Cory C. (October 13, 2009). "Global patterns in belowground communities" (in en). Ecology Letters 12 (11): 1238–1249. doi:10.1111/j.1461-0248.2009.01360.x. PMID 19674041. https://pubmed.ncbi.nlm.nih.gov/19674041/.
43. ↑ Edwards, Clive A.; Normal, Arancon Q. (April 27, 2022). "Earthworm Physiology". Biology and Ecology of Earthworms (4 ed.). New York, NY: Springer. pp. 33–54. ISBN 978-0-387-74943-3.
44. ↑ Tuma, Jiri; Eggleton, Paul; Fayle, Tom M. (25 Dec 2019). "Ant‐termite interactions: an important but under‐explored ecological linkage" (in en). Biological Reviews 95 (3): 555–572. doi:10.1111/brv.12577. ISSN 1464-7931. PMID 31876057. https://onlinelibrary.wiley.com/doi/10.1111/brv.12577.
45. ↑ 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
46. ↑ 46.0 46.1 van den Hoogen, Johan (8 August 2019). "Soil nematode abundance and functional group composition at a global scale". Nature 572. doi:10.1038/s41586-019-1418-6.
47. ↑ 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)
48. ↑ 48.0 48.1 "Whaling and Deep-Sea Biodiversity". Conservation Biology 10 (2): 653–654. 1996. doi:10.1046/j.1523-1739.1996.10020653.x.
49. ↑ Fisheries, NOAA (2022-05-03). "Fun Facts About Fascinating Fish | NOAA Fisheries" (in en). https://www.fisheries.noaa.gov/national/outreach-and-education/fun-facts-about-fascinating-fish.
50. ↑ Johnson, Brett M.; Pate, William M.; Hansen, Adam G. (2017). "Energy Density and Dry Matter Content in Fish: New Observations and an Evaluation of Some Empirical Models". Transactions of the American Fisheries Society 146 (6): 1262–1278. doi:10.1080/00028487.2017.1360392.
51. ↑ 51.0 51.1 51.2 "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.
52. ↑ "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.
53. ↑ Alexander, David E. (1 May 1999). Encyclopedia of Environmental Science. Springer. ISBN 978-0-412-74050-3.
54. ↑ "What are wetlands?". https://www.ramsar.org/sites/default/files/documents/library/info2007-01-e.pdf.
55. ↑ 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.
56. ↑ Mark Spalding, Corinna Ravilious, and Edmund Green. 2001. World Atlas of Coral Reefs. Berkeley, California: University of California Press and UNEP/WCMC.
57. ↑ 57.0 57.1 57.2 57.3 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.
58. ↑ "Tundra - Biomes - WWF". https://www.worldwildlife.org/biomes/tundra.
59. ↑ "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

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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

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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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