Global nutrient transport in a world of giants
Open accessAbstract
Significance: Animals play an important role in the transport of nutrients, but this role has diminished because many of the largest animals have gone extinct or experienced massive population declines. Here, we quantify the movement of nutrients by animals in the land, sea, rivers, and air both now and prior to their widespread reductions. The capacity to move nutrients away from hotspots decreased to 6% of past values across land and ocean. The vertical movement of phosphorus (P) by marine mammals was reduced by 77% and movement of P from sea to land by seabirds and anadromous fish was reduced by 96%, effectively disrupting an efficient nutrient distribution pump that once existed from the deep sea to the continental interiors.
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- Findingown data§ Results · p. 3
Current marine mammal vertical phosphorus movement is about 75 million kg P/yr, with a range of 54 to 110 million.
“they now move ∼ 75 million (54 -110 million; SI Appendix , Table 2) kg of P per year”
Quote in the paperSubstantial claim - Findingown data§ Global nutrient transport in a world of giants · p. 2
Ocean nutrient distribution capacity by great whales is currently about 5% of historic values.
“We estimate that the capacity of animals to move nutrients away from concentration patches has decreased to about 8% of the preextinction value on land and about 5% of historic values in oceans.”
- Also stated in: § Results · p. 4 · plain
“Current mean, km 2 · y - 1 | 1.6e4 | 3.2e4 | 6.67e4 | 467 | 1.13e4 | 0.42e4 | 0.17e4 | 0.22e5 |”
Quote in the paperSubstantial claim - Also stated in: § Results · p. 4 · plain
- Findingown data§ Global nutrient transport in a world of giants · p. 2
Past marine mammal vertical phosphorus movement was about 340 million kg P/yr, with a range of 260 to 430 million.
“For phosphorus (P), a key nutrient, upward movement in the ocean by marine mammals is about 23% of its former capacity (previously about 340 million kg of P per year).”
- Also stated in: § Results · p. 3 · plain
“they moved a global total of ∼ 340 million (260 -430 million; SI Appendix , Table 2) kg of P per year from the depth to the surface waters before widespread hunting”
Quote in the paperSubstantial claim - Also stated in: § Results · p. 3 · plain
- Findingown data§ Results · p. 3
Current global mean terrestrial nutrient distribution capacity is 16,000 km2/yr.
“currently 16,000 km 2 · y -1”
Quote in the paperSubstantial claim - Findingown data§ Global nutrient transport in a world of giants · p. 2
Marine mammal vertical phosphorus movement is now about 23% of its former capacity.
“For phosphorus (P), a key nutrient, upward movement in the ocean by marine mammals is about 23% of its former capacity (previously about 340 million kg of P per year).”
- Also stated in: § Results · p. 3 · plain
“representing a decrease to 23% of original capacity (Fig. 2 D -F and Table 1).”
Quote in the paperSubstantial claim - Also stated in: § Results · p. 3 · plain
- Findingnone given§ Global nutrient transport in a world of giants · p. 2
Past sea-to-land phosphorus movement by seabirds and anadromous fish totalled about 150 million kg P/yr globally.
“Movements by seabirds and anadromous fish provide important transfer of nutrients from the sea to land, totalling ∼ 150 million kg of P per year globally in the past”
Quote in the paperSubstantial claim- The authors state this as an assumption: 20% of seabird guano produced reaches coastal land, with a range of 5% to 35%.
- The authors state this as an assumption: Seabird population calculations are representative of past, not current, populations because they are based on theoretical population densities.
- Findingown data§ Results · p. 3
Global mean terrestrial nutrient distribution capacity before the late-Quaternary extinctions averaged 180,000 km2/yr.
“averaged 180,000 km 2 · y -1 on the land surface”
Quote in the paperSubstantial claim - Findingreasoning§ Global nutrient transport in a world of giants · p. 2
Current sea-to-land phosphorus movement by seabirds and anadromous fish is less than 4% of the past value.
“a transfer that has declined to less than 4% of this value as a result of the decimation of seabird colonies and anadromous fish populations.”
- Also stated in: § Significance · p. 2 · plain
“movement of P from sea to land by seabirds and anadromous fish was reduced by 96%”
Quote in the paperSubstantial claim- The authors state this as an assumption: 20% of seabird guano produced reaches coastal land, with a range of 5% to 35%.
- Also stated in: § Significance · p. 2 · plain
- Findingreasoning§ Significance · p. 2
The capacity to move nutrients away from hotspots decreased to 6% of past values across land and ocean.
“The capacity to move nutrients away from hotspots decreased to 6% of past values across land and ocean.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Vertical transport capacity in the North Atlantic is 28% of its historical value.
“and the North Atlantic (28%)”
Quote in the paperNot a substantial claim - Findingreasoning§ Discussion · p. 4
Currently, with the largest animals in South America weighing only up to about 300 kg, continental nutrient distribution capacity has dropped to about 0.01% of its original value in some regions.
“However, currently, with the largest animals in South America only weighing up to ∼ 300 kg, continental nutrient distribution capacity has dropped to ∼ 0.01% of its original value in some regions.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Every continent outside Africa is at less than 5% of its original nutrient distribution capacity.
“Every continent outside Africa (Africa is at 46% of its lateQuaternary value) is at less than 5% of the original value”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 4
Historically, anadromous fish may have moved at least an order of magnitude more phosphorus from ocean to land than seabirds, with an estimate of 140 million (71-430 million) km^2 y^-1 versus 6.3 million (1.5-16 million) km^2 y^-1 for seabirds.
“Using range maps for 42 of these species, and an additional 47 closely related species as proxies for the missing range maps, we estimate that, historically, anadromous fish may have moved at least an order of magnitude more P from ocean to land [140 million (71 -430 million) km 2 · y -1 ; SI Appendix , Table 2] than seabirds [6.3 million (1.5 -16 million) km 2 · y -1 ; SI Appendix , Table 2]”
- Also stated in: § Discussion · p. 5 · hedged
“Possibly, a more important form of sea-to-land nutrient transport comes from the migratory behavior of anadromous fish, which we estimate to bring at least an order of magnitude more nutrients from oceans to land than do seabirds.”
Quote in the paperSubstantial claim - Also stated in: § Discussion · p. 5 · hedged
- Findingown data§ Results · p. 3
Great whale lateral diffusion capacity in the North Atlantic is 14% of its historical value.
“and the North Atlantic (14%)”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Africa is at 46% of its late-Quaternary nutrient distribution capacity.
“Africa is at 46% of its lateQuaternary value”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Great whale lateral diffusion capacity in the Southern Ocean is 2% of its historical value.
“estimated that the capacity in the Southern Ocean is 2% of its historical value”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Without historical range reduction of large species, excluding extinctions, lateral nutrient flux capacity would be 37% higher than today's baseline.
“we estimate that without the range reduction of large species (excluding all extinctions) the capacity would be 37% higher compared with today ' s baseline.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
In the past, marine mammals could have increased surface phosphorus concentrations by up to 1% per year in the Southern Ocean.
“in the past, marine mammals could have increased surface concentrations by up to 1% per year in the Southern Ocean”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Mean nutrient diffusion capacity is larger for great whales than for terrestrial animals at natural capacity, at 640,000 km2/yr versus 180,000 km2/yr.
“Mean nutrient diffusion capacity is larger for the great whales than for terrestrial animals at natural capacity (640,000 km 2 · y -1 for great whales vs. 180,000 km 2 · y -1 for terrestrial mammals).”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Great whale lateral diffusion capacity in the North Pacific is 10% of its historical value.
“with slightly higher values in the North Pacific (10%)”
Quote in the paperSubstantial claim - Findingnone given§ Discussion · p. 4
The decimation of terrestrial megafauna and whales reduced the ability of animals to distribute nutrients away from nutrient-abundant regions to about 6% of global natural capacity.
“We estimate that the decimation of terrestrial megafauna and whales has reduced the ability of animals to distribute nutrients away from regions of nutrient abundance to ∼ 6%of global natural capacity.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 4
Anadromous fish phosphorus flux values are not evenly distributed, with higher values in the Northern Hemisphere and at high latitudes than in tropical latitudes.
“These values are not evenly distributed, and there are much higher values in the Northern Hemisphere and at high latitudes than in more tropical latitudes.”
Quote in the paperNot a substantial claim - Findingreasoning§ Discussion · p. 4
This large number of megaherbivores gave southern South America, before the extinctions, the largest estimated present natural lateral diffusion capacity of anywhere in the world.
“This large number of megaherbivores gave it, before the extinctions, the largest estimated present natural lateral diffusion capacity of anywhere in the world.”
Quote in the paperSubstantial claim - Findingreasoning§ Discussion · p. 4
On land, disparities in animal sizes and total numbers led to differences in nutrient distribution capacity before and after extinctions because extinctions were regionally uneven.
“On land, large disparities in animal sizes and total animal numbers led to differences in nutrient distribution capacity both before and following the extinctions due to regional disparities in extinctions.”
Quote in the paperSubstantial claim - Findingown data§ Results · p. 3
Vertical transport capacity in the North Pacific is 34% of its historical value.
“but there are higher values in the North Pacific (34%)”
Quote in the paperNot a substantial claim - Findingown data§ Results · p. 3
Vertical transport capacity in the Southern Ocean is now about 16% of its historical value.
“Vertical transport capacity in the Southern Ocean is now ∼ 16% of its historical value”
Quote in the paperSubstantial claim - Findingown data§ Discussion · p. 5
Seabirds can increase phosphorus concentrations in coastal environments globally by about 6 million kg y^-1 through guano deposition.
“Here, we have estimated that seabirds can increase P concentrations in coastal environments globally by ∼ 6 million kg · y -1 through the deposition of guano.”
Quote in the paperSubstantial claim- The authors state this as an assumption: Twenty percent of guano arrives on land.
- Findingown data§ Results · p. 3
South America has the largest change, at about 1% of its original nutrient distribution capacity.
“with the largest change in South America ( ∼ 1% of the original value; Table 1).”
Quote in the paperNot a substantial claim - Findingcitation§ Discussion · p. 5
The total past flux of phosphorus from sea to land by anadromous fish and seabirds (146 million kg P per year) is much less than the total phosphorus consumed by humans for fertilizers each year (48,500 million kg phosphoric acid as P2O5 in 2010, growing at 1.9% per year).
“We estimate the total flux of P from sea to land by anadromous fish and seabirds in the past (146 million kg of P per year) is still much less than the total P consumed by humans for fertilizers each year [48,500 million kg of phosphoric acid (as P2O5) in 2010 and growing at 1.9% per year (39)].”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [39] Food and Agricultural Organization of the United Nations (2011) Current world fertilizer trends and outlook to 2015. Available at www.fao.org/3/a-av252e.pdf. Accessed September 4, 2015.
Not checked in the free run.
- Premisecitation§ Global nutrient transport in a world of giants · p. 2
Around 150 mammal megafaunal species, defined as at least 44 kg body mass, went extinct in the late Pleistocene and early Holocene.
“around 150 mammal megafaunal (here, defined as ≥ 44 kg of body mass) species going extinct in the late Pleistocene and early Holocene (1, 2).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation§ Significance · p. 2
Animals play an important role in transporting nutrients.
“Animals play an important role in the transport of nutrients, but this role has diminished because many of the largest animals have gone extinct or experienced massive population declines.”
Quote in the paperSubstantial claim0 of 3 sources checked - Premisecitation§ Global nutrient transport in a world of giants · p. 2
Whale densities might have declined between 66% and 99%.
“whale densities might have declined between 66% and 99% (4 -6).”
Quote in the paperSubstantial claim0 of 3 sources checked - Interpretationown data§ Significance · p. 2
The role of animals in nutrient transport has diminished because many of the largest animals went extinct or experienced massive population declines.
“Animals play an important role in the transport of nutrients, but this role has diminished because many of the largest animals have gone extinct or experienced massive population declines.”
Quote in the paperSubstantial claim - Premisecitation§ Significance · p. 3
The distribution of nutrients away from a concentration gradient is size-dependent, with larger animals disproportionately more important than smaller animals.
“the distribution of nutrients away from a concentration gradient is size-dependent, with larger animals having disproportionally greater importance to this flow of nutrients than smaller animals (20, 21).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation§ Global nutrient transport in a world of giants · p. 2
Animal digestion accelerates cycling of nutrients from recalcitrant plant matter to labile forms in excreta after herbivore consumption on land.
“Animal digestion accelerates cycling of nutrients from more recalcitrant forms in decomposing plant matter to more labile forms in excreta after (wild or domestic) herbivore consumption on land (7).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [7] source
Not checked in the free run.
- Premisecitation§ Global nutrient transport in a world of giants · p. 2
Southern Hemisphere blue whales were reduced to 1% of their historical numbers as a result of commercial whaling.
“in the Southern Hemisphere, blue whales ( Balaenoptera musculus ) have been reduced to 1% of their historical numbers as a result of commercial whaling (4).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [4]
Not checked in the free run.
- Premisecitation§ Global nutrient transport in a world of giants · p. 2
At present, large herbivores enhance nutrient cycling in the grazing lawns of the Serengeti.
“Similarly, at present times, large herbivores enhance nutrient cycling in the grazing lawns of the Serengeti (9).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [9] source
Not checked in the free run.
- Interpretationcitation§ Discussion · p. 5
This nutrient flux has almost certainly decreased through time as seabird populations have decreased or gone extinct, often due to invasive mammal predators.
“This flux of nutrients has almost certainly decreased through time as seabird populations have decreased [27% of seabirds are classified as threatened (22)] or gone extinct (e.g., the great auk, Pinguinus impennis ) often due to, for instance, invasive mammal predators decimating seabird colonies (36).”
Quote in the paperNot a substantial claim0 of 2 sources checked - Premisecitation§ Discussion · p. 4
In some regions, exotic and mainly domesticated ungulates may have partially taken over the nutrient distribution role.
“However, in some regions, exotic and mainly domesticated ungulates may have partially taken over this role (33).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [33] Pires MM, et al. (2014) Reconstructing past ecological networks: The reconfiguration of seed-dispersal interactions after megafaunal extinction. Oecologia 175(4):1247 -1256. source
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- Premisecitation§ Results · p. 4
There are 110 species of anadromous fish that migrate from oceans to rivers to breed and eventually die.
“It is estimated that there are 110 species of anadromous fish, such as salmon, that migrate from oceans to rivers to breed and eventually die ( SI Appendix , Table 5) (31).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [31] McDowall RM (1988) Diadromy in Fishes: Migrations Between Freshwater and Marine Environments (Croom Helm, London).
Not checked in the free run.
- Interpretationown data§ Results · p. 3
Great whales might have once transported nutrients away from concentration gradients more efficiently than terrestrial mammals.
“great whales might have once transported nutrients away from concentration gradients more efficiently than terrestrial mammals.”
Quote in the paperSubstantial claim - Premisecitation§ Discussion · p. 4
Regional models found that phosphorus transfer away from Amazonian floodplains may have dropped by more than 50% after the extinction of Amazonian megafauna.
“Regional models found that the transfer of P away from the Amazonian floodplains may have dropped by more than 50% following the extinction of the Amazonian megafauna (20, 21).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation§ Discussion · p. 4
Previous experimental studies found that animals move significant quantities of nutrients across concentration gradients even when they do not necessarily move dung from fertile to nonfertile areas.
“Previous experimental studies, however, have found that animals move significant quantities of nutrients across concentration gradients despite not necessarily moving dung from fertile to nonfertile areas (11, 14, 19).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation§ Discussion · p. 5
A much larger share of primary production in oceans is consumed compared with terrestrial primary producers.
“Furthermore, a much larger share of primary production in oceans (algae) is consumed compared with terrestrial primary producers (35).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [35] Cyr H, Pace ML (1993) Magnitude and patterns of herbivory in aquatic and terrestrial ecosystems. Nature 361(6408):148 -150. source
Not checked in the free run.
- Premisecitation§ Discussion · p. 5
Whales and their prey may help retain limiting nutrients (N, P, and Fe) in the surface layer and release them slowly into the water.
“Also, whales and their prey may help in retaining limiting nutrients (N, P, and Fe) in the surface layer and releasing these nutrients slowly into the water (18).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [18] source
Not checked in the free run.
- Premisecitation§ Discussion · p. 5
Marine mammals have experienced broad population reductions due to widespread hunting over the past few hundred years.
“Marine mammals have seen broad population reductions due to widespread hunting over the past few hundred years (34).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [34] Branch TA, et al. (2007) Past and present distribution, densities and movements of blue whales Balaenoptera musculus in the Southern Hemisphere and northern Indian Ocean. Mammal Rev 37(2):116 -175. source
Not checked in the free run.
- Interpretationreasoning§ Discussion · p. 5
Limiting nutrients should be converted to primary producer biomass more quickly in oceans than on land.
“and due to this faster turnover time, nutrients, especially limiting nutrients, should be converted to primary producer biomass more quickly in the oceans.”
Quote in the paperSubstantial claim- Rests on an assumption: Aquatic algae conduct most of the ocean's photosynthesis and have a much faster turnover time than land plants due to their often single-cellular nature.
- Interpretationreasoning§ Results · p. 3
This could result in considerable stock changes in surface phosphorus over time.
“which could result in considerable stock changes in surface P over time.”
Quote in the paperSubstantial claim - Interpretationreasoning§ Discussion · p. 5
Seabirds may act as a link connecting nutrient concentrations in the oceans with nutrient concentrations on land.
“Seabirds may act as a link connecting nutrient concentrations in the oceans with nutrient concentrations on the land.”
Quote in the paperSubstantial claim - Interpretationnone given§ Discussion · p. 5
Anadromous fish are especially important vectors because they travel much further inland than seabirds.
“Anadromous fish seem to be especially important vectors of nutrients because they travel much further inland than seabirds (Fig. 3).”
Quote in the paperSubstantial claim - Interpretationreasoning§ Discussion · p. 6
Before the widespread extinction of megafauna and hunting of whales, an interlinked system for recycling nutrients may have existed, with nutrients flowing against entropy from ocean depths to continental interiors.
“Before the widespread extinction of megafauna and hunting of whales, an interlinked system for the recycling of nutrients may have existed, where nutrients flowed against entropy from the ocean depths to continental interiors (Fig. 4).”
Quote in the paperSubstantial claim - Interpretationnone given§ Discussion · p. 6
In this system, marine mammals moved nutrients vertically to the surface, increasing productivity.
“Marine mammals moved nutrients vertically to the surface, increasing productivity.”
Quote in the paperNot a substantial claim - Interpretationown data§ Discussion · p. 6
A world teeming with large animals may have had an efficient system of redistributing phosphorus.
“We have shown that a world teeming with large animals may have had an efficient system of redistributing P.”
Quote in the paperSubstantial claim - Premisecitation§ Discussion · p. 6
These processes can regulate atmospheric CO2 levels through stimulation of new primary production and consequent export of carbon to the deep ocean.
“These processes can regulate atmospheric CO2 levels through the stimulation of new primary production and consequent export of carbon to the deep ocean (16, 44).”
Quote in the paperSubstantial claim0 of 2 sources checked - Interpretationown data§ Discussion · p. 6
The paper has demonstrated the plausibility of an animal-mediated chain of nutrient transfer connecting the deep ocean to continental interiors.
“Although the numbers we have calculated in this paper are exploratory (we explore this uncertainty in SI Appendix , Tables 1 and 2) and subject to further research and quantification, we have demonstrated the plausibility of an animal-mediated chain of nutrient transfer that connects the deep ocean to the continental interiors.”
Quote in the paperSubstantial claim - Premisecitation§ Discussion · p. 5
Isotopic evidence indicates that significant quantities of ocean-derived nutrients from anadromous fish enter terrestrial ecosystems.
“However, isotopic evidence indicates that significant quantities of ocean-derived nutrients from anadromous fish do enter terrestrial ecosystems (11).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [11]
Not checked in the free run.
- Premisecitation§ Discussion · p. 6
At current rates of use, known global stocks of phosphate rock are estimated to run out in as little as 50 years.
“At current rates of use, the known global stocks of phosphate rock are estimated to run out in as little as 50 y (45).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [45] Gilbert N (2009) Environment: The disappearing nutrient. Nature 461(7265):716 -718. source
Not checked in the free run.
- Interpretationreasoning§ Discussion · p. 6
Each step in the proposed system is potentially a nonlinear positive feedback of increasing productivity.
“Each step is potentially a nonlinear positive feedback of increasing productivity.”
Quote in the paperSubstantial claim - Interpretationreasoning§ Discussion · p. 6
On longer time scales, coastal seabird island nutrient hotspots may become exposed to terrestrial fauna and spread toward continental interiors.
“Finally, on longer time scales, coastal seabird island nutrient hotspots may become exposed to terrestrial fauna and spread toward continental interiors.”
Quote in the paperSubstantial claim- Rests on an assumption: Guano is generally deposited on steep cliffs or offshore islands that are largely inaccessible to most terrestrial animals.
- Rests on an assumption: Most seabird islands are on continental shelves that become connected to continents during interglacial periods.
- May combine two claims. 26
- Classified as probably not a substantial claim. 5
May combine two claims.
Findingown data§ Global nutrient transport in a world of giants · p. 2Global mean terrestrial nutrient distribution capacity is currently about 8% of its pre-extinction value, with a past mean of 180,000 km2/yr and a current mean of 16,000 km2/yr.
“We estimate that the capacity of animals to move nutrients away from concentration patches has decreased to about 8% of the preextinction value on land and about 5% of historic values in oceans.”
- Also stated in: § Results · p. 3 · hedged
“We estimate that the global mean nutrient distribution capacity before the late-Quaternary extinctions averaged 180,000 km 2 · y -1 on the land surface and that it is currently 16,000 km 2 · y -1 , ∼ 8% of its former value (Table 1; detailed methodology is provided in Methods and SI Appendix ).”
Quote in the paperSubstantial claim- Also stated in: § Results · p. 3 · hedged
May combine two claims.
Findingown data§ Results · p. 4Global average sea-to-land phosphorus movement in coastal regions is 0.19 ± 0.15 kg P km^-2 y^-1, varying by an order of magnitude across the planet with peaks in the Southern Hemisphere.
“Therefore global averaged sea-to-land movement of P is 0.19 ± 0.15 kg of P km -2 · y -1 in coastal regions, but varies by an order of magnitude throughout the planet, with peaks in the Southern Hemisphere (Fig. 3 and Table 1).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ Results · p. 4Current anadromous fish phosphorus flux has decreased to about 4% or less of historic values, corresponding to 5.6 million km^2 y^-1.
“but this estimated value has decreased to ∼ 4% (5.6 million km 2 · y -1 ) of the original value.”
- Also stated in: § Discussion · p. 5 · hedged
“However, they have also experienced drastic population losses (25 -27), and we estimate that the current nutrient flux is less than 4% of historic values, before overfishing and habitat modification, such as damming of rivers.”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [25] Gresh T, Lichatowich J, Schoonmaker P (2000) An estimation of historic and current levels of salmon production in the Northeast Pacific ecosystem: Evidence of a nutrient deficit in the freshwater systems of the Pacific Northwest. Fisheries (Bethesda, Md) 25(1):15 -21.
Not checked in the free run.
- cites [26] de Groot SJ (2002) A review of the past and present status of anadromous fish species in the Netherlands: Is restocking the Rhine feasible? Ecological restoration of aquatic and semi-aquatic ecosystems in the Netherlands (NW Europe). Developments in Hydrobiology 166:205 -218. source
Not checked in the free run.
- cites [27] Limburg KE, Waldman JR (2009) Dramatic declines in North Atlantic diadromous fishes. Bioscience 59(11):955 -965. source
Not checked in the free run.
- Also stated in: § Discussion · p. 5 · hedged
May combine two claims.
Findingown data§ Results · p. 3Before the extinctions, nutrient distribution capacity was more evenly spread than currently, with most current capacity only in Africa.
“Before the extinctions, nutrient distribution capacity was much more evenly spread than it is currently, with most of the current capacity only in Africa, where extensive megafauna remain.”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ Results · p. 4The global flux of phosphorus from sea to land by seabirds is 6.3 million (1.5-16 million) kg km^-2 y^-1, with almost half moving onto the Eurasian landmass.
“We calculate a global flux of 6.3 million (1.5 -16 million; SI Appendix , Table 2) kg km -2 · y -1 of P from sea to land by the seabirds, with almost half moving onto the Eurasian landmass (Table 1).”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ Results · p. 3In parts of Africa such as Kruger National Park, nutrient distribution capacity remains close to 100% of Late Pleistocene values, while southern South America is below 0.01% of previous values.
“in parts of Africa, such as Kruger National Park in South Africa, capacity is still close to 100% of what it once was in the Late Pleistocene, whereas other regions, such as southern South America, are at less than 0.01% of previous values”
Quote in the paperSubstantial claimMay combine two claims.
Findingown data§ Results · p. 4When averaged by coastal continental area, phosphorus movement is highest in Oceania at 0.31 ± 0.23 kg P km^-2 y^-1 and lower in North America at 0.16 ± 0.12 kg P km^-2 y^-1.
“When averaged by coastal continental area, we find a maximum in Oceania, with 0.31 ± 0.23 kg of P km -2 · y -1 , and lower values in North America, with 0.16 ± 0.12 kg of P km 2 · y -1 .”
Quote in the paperNot a substantial claimMay combine two claims.
Findingreasoning§ Discussion · p. 4South America once had the largest nutrient distribution capacity, but after the Late-Pleistocene extinctions it experienced the largest drop, to about 1% of its original continent-wide capacity.
“For instance, South America once had the largest nutrient distribution capacity, but following the Late-Pleistocene extinctions, it experienced the largest drop, to ∼ 1% of its original continent-wide capacity.”
Quote in the paperSubstantial claimMay combine two claims.
Premisecitation§ Significance · p. 327% of all seabirds are classified as threatened, and up to 75% of albatross species are considered threatened or endangered.
“27% of all seabirds are classified as threatened (critically endangered, endangered, or vulnerable), and the largest of all seabirds, the albatross, is the most endangered, with up to 75% of albatross species considered threatened or endangered (22 -24)”
Quote in the paperSubstantial claim0 of 3 sources checkedMay combine two claims.
Premisecitation§ Significance · p. 3Anadromous fish populations have declined to less than 10% of their historical numbers in the Pacific Northwest and in both the northeastern and northwestern Atlantic.
“populations of anadromous fish have declined to less than 10% of their historical numbers in the Pacific Northwest (25) and both the northeastern and northwestern Atlantic (26, 27).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites [25] Gresh T, Lichatowich J, Schoonmaker P (2000) An estimation of historic and current levels of salmon production in the Northeast Pacific ecosystem: Evidence of a nutrient deficit in the freshwater systems of the Pacific Northwest. Fisheries (Bethesda, Md) 25(1):15 -21.
Not checked in the free run.
- cites [26] de Groot SJ (2002) A review of the past and present status of anadromous fish species in the Netherlands: Is restocking the Rhine feasible? Ecological restoration of aquatic and semi-aquatic ecosystems in the Netherlands (NW Europe). Developments in Hydrobiology 166:205 -218. source
Not checked in the free run.
- cites [27] Limburg KE, Waldman JR (2009) Dramatic declines in North Atlantic diadromous fishes. Bioscience 59(11):955 -965. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing).
Interpretationcitation§ Discussion · p. 5In the past, scavenging birds such as condors may also have acted as vectors of nutrients from the sea to land.
“In the past, scavenging birds, such as condors, may also have acted as vectors of nutrients from the sea to the land.”
Quote in the paperNot a substantial claimProbably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductown data§ Results · p. 4Each calculated value has many uncertainties, which are explored in a sensitivity study.
“Each value has many uncertainties involved in its calculation, which we explore in a sensitivity study ( SI Appendix , Tables 1 and 2).”
Quote in the paperNot a substantial claimMay combine two claims.
Premisecitation§ Discussion · p. 5Once nutrients drop below the photic zone into deep ocean sediments, they are generally considered lost to surface biota, and only tectonic movements and limited upwelling recycle them.
“This ability to spread nutrients vertically may be especially important, because once nutrients drop below the photic zone and into the deep ocean sediments, they are generally considered to be lost to the surface biota, and only tectonic movements and limited regions where water is uplifted, will further recycle them (16).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [16] source
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ Discussion · p. 5Such population decreases reduced lateral distribution capacity and perhaps reduced vertical distribution, allowing more nutrients to drop below the photic zone.
“Such population decreases reduced the lateral distribution capacity and perhaps reduced the vertical distribution, allowing more nutrients to drop below the photic zone.”
Quote in the paperNot a substantial claimMay combine two claims.
Premisecitation§ Discussion · p. 4Southern South America once had the largest number of megaherbivores over 1,000 kg (n = 15), all of which went extinct.
“For instance, southern South America once had the largest number of megaherbivores ( > 1,000 kg, n = 15), all of which went extinct (32).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [32] Owen-Smith N (2013) Contrasts in the large herbivore faunas of the southern continents in the late Pleistocene and the ecological implications for human origins. J Biogeogr 40(7):1215 -1224. source
Not checked in the free run.
May combine two claims.
Interpretationcitation§ Discussion · p. 5This flux has certainly greatly decreased as both marine mammals and large scavengers have seen their numbers drop substantially.
“This flux has certainly greatly decreased as both marine mammals and large scavengers have seen their numbers drop substantially (38).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [38] Tyrberg T (2008) The Late Pleistocene continental avian extinction -An evaluation of the fossil evidence. Oryctos 7:249 -269.
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ Discussion · p. 5Nutrients transported by whales, or as a consequence of whale activity, should be assimilated more rapidly and contribute to system productivity more directly than on land.
“The nutrients transported by whales, or as a consequence of whale activity, should be assimilated more rapidly, and contribute to system productivity more directly than on land.”
Quote in the paperSubstantial claimMay combine two claims.
Study design and conductcitation§ Results · p. 4These estimates are based on theoretical population densities and are probably closer to theoretical historical values than to actual values today.
“These estimates are calculated based on theoretical population densities (30), and it is probably closer to theoretical historical values than to actual values today.”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [30] Juanes F (1986) Population-density and body size in birds. Am Nat 128(6):921 -929. source
Not checked in the free run.
May combine two claims.
Interpretationcitation§ Discussion · p. 6Redistribution of phosphorus from areas of excess to phosphorus-poor soils may simultaneously boost global crop production and reduce eutrophication.
“Therefore, a redistribution of P from areas where it is currently found in excess to areas where the soil is naturally P-poor may simultaneously boost global crop production and reduce eutrophication (47).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [47] Steffen W, et al. (2015) Sustainability. Planetary boundaries: Guiding human development on a changing planet. Science 347(6223):1259855. source
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ Discussion · p. 6Although the fluxes can be calculated separately, the net increase in global productivity is likely more than the sum of the four systems together.
“Given these and many other connections, it is clear that although we can calculate each of these fluxes (marine mammals, seabirds, anadromous fish, and megafauna) separately, the net increase in global productivity is likely more than the sum that we have calculated of the four systems together.”
Quote in the paperSubstantial claimMay combine two claims.
Interpretationreasoning§ Discussion · p. 6Restoration of whale populations could increase productivity by transferring nutrients to oligotrophic areas and enhancing Fe and N at the ocean surface.
“The restoration of whale populations could increase productivity by transferring nutrients to oligotrophic areas and enhancing Fe and N at the ocean surface.”
Quote in the paperNot a substantial claimClassified as probably not a substantial claim. May combine two claims.
Premisecitation§ Methods · p. 7Lateral nutrient distribution capacity was mathematically formulated and found to be strongly size-dependent in two previous papers, and this framework is reformulated in the SI Appendix.
“Lateral nutrient distribution capacity was mathematically formulated and found to be strongly size-dependent in two previous papers (20, 21), and this mathematical framework is reformulated in SI Appendix .”
Quote in the paperSubstantial claim0 of 2 sources checkedMay combine two claims.
Premisecitation§ Discussion · p. 6Phosphorus is not well distributed, causes eutrophication in some areas, and deficits affect almost 30% of global cropland area.
“P is not well distributed, and it causes eutrophication in some areas, whereas P deficits still affect almost 30% of the global cropland area (46, 47).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites [46] MacDonald GK, Bennett EM, Potter PA, Ramankutty N (2011) Agronomic phosphorus imbalances across the world ' s croplands. Proc Natl Acad Sci USA 108(7):3086 -3091. source
Not checked in the free run.
- cites [47] Steffen W, et al. (2015) Sustainability. Planetary boundaries: Guiding human development on a changing planet. Science 347(6223):1259855. source
Not checked in the free run.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Interpretationcitation§ Discussion · p. 5The loss of nutrients to these ecosystems from historic highs may have affected the entire ecosystem, including the fish themselves, and contributed to the downward spiral of salmonid abundance and diversity.
“This loss of nutrients to these ecosystems from historic highs may have affected the entire ecosystem, including the fish themselves, and ' contributed to the downward spiral of salmonid abundance and diversity in general ' (25).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites [25] Gresh T, Lichatowich J, Schoonmaker P (2000) An estimation of historic and current levels of salmon production in the Northeast Pacific ecosystem: Evidence of a nutrient deficit in the freshwater systems of the Pacific Northwest. Fisheries (Bethesda, Md) 25(1):15 -21.
Not checked in the free run.
May combine two claims.
Interpretationreasoning§ Discussion · p. 6Although the biomass of domestic animals is now much greater than the total biomass of extinct large terrestrial animals, it is unlikely that domestic animals distribute nutrients in the manner that existed in the world of giant megafauna.
“Although the biomass of domestic animals is now much greater than the total biomass of the extinct large terrestrial animals (42), there are two reasons why it is unlikely that they distribute nutrients in the manner that we believe existed in the world of giant megafauna.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites [42] Barnosky AD (2008) Colloquium paper: Megafauna biomass tradeoff as a driver of Quaternary and future extinctions. Proc Natl Acad Sci USA 105(Suppl 1):11543 -11548. source
Not checked in the free run.
- Rests on an assumption: Most domestic animals in the industrialized world are fenced in or indoors, and fences constrain movement.
- Rests on an assumption: Most pastures contain only a single animal type, such as cattle.
- Rests on an assumption: A single species is more likely to have consistent behavior, eating in one place and defecating in another, concentrating nutrients rather than dispersing them.
- Rests on an assumption: Multiple species have different eating and defecating patterns and are more likely to diffuse nutrients across gradients.
Probably not a substantial claim: may not pass S3 (load-bearing). May combine two claims.
Study design and conductreasoning§ Discussion · p. 5It is uncertain what quantity of nutrients transported inland by fish arrives onto land, but it is clearly a function of river size, distance transported inland, and consumption by scavengers and predators.
“It is uncertain what quantity of nutrients transported inland by the fish arrives onto terra firma , but it is clearly a function of river size, distance transported inland, and consumption of the fish by scavengers and predators.”
Quote in the paperNot a substantial claimMay combine two claims.
Interpretationreasoning§ Discussion · p. 6Increased surface productivity increased available food for seabirds and anadromous fish, potentially increasing the flux of nutrients from sea to land.
“This increased surface productivity increased available food for seabirds and anadromous fish, potentially increasing the flux of nutrients from sea to land.”
Quote in the paperSubstantial claim
The authors state this as an assumption.
No previous study has attempted a global-scale estimate of how nutrient distribution changed from before human-caused extinctions and exploitation to today across oceans, air, rivers, and land.
“There have been many individual site-level studies showing the importance of animals in distributing nutrients, but as far as we are aware, no previous study has attempted to estimate at a global scale how this distribution has changed from the time before humancaused extinctions and exploitation up to today in the oceans, air, rivers, and land.”
The paper relies on this without support.
Phosphorus serves as a proxy for other limiting elements such as nitrogen and iron.
“We calculate nutrient fluxes caused by animals in terms of the frequently limiting nutrient, P, which serves as a proxy for other limiting elements, such as N and Fe.”
The authors state this as an assumption.
20% of seabird guano produced reaches coastal land, with a range of 5% to 35%.
“assume 20% (5 to 35%) of guano reaches land”
“We assume that 20% (ranging between 5% and 35%) of seabird guano produced arrives on coastal land.”
The authors state this as an assumption.
Seabird population calculations are representative of past, not current, populations because they are based on theoretical population densities.
“We assume our calculations for seabird populations are representative of past, not current, populations because they are based on theoretical population densities (43).”
The paper relies on this without support.
Guano is generally deposited on steep cliffs or offshore islands that are largely inaccessible to most terrestrial animals.
“Guano is generally deposited on steep cliffs or offshore islands generally inaccessible to most terrestrial animals.”
The paper relies on this without support.
Most seabird islands are on continental shelves that become connected to continents during interglacial periods.
“For instance, most seabird islands are on the continental shelves, which become connected to the continent during interglacial periods, thus exposing the nutrient hotspots to terrestrial animals.”
The paper relies on this without support.
Most domestic animals in the industrialized world are fenced in or indoors, and fences constrain movement.
“First, most domestic animals in the industrialized world are fenced in (or indoors), and fences constrain movement, thus stopping the diffusion of nutrients (fences are less common in the nonindustrialized world, although the animals may be fenced in at night).”
The paper relies on this without support.
Most pastures contain only a single animal type, such as cattle.
“Second, most pastures have only a single animal type, such as cattle.”
The paper relies on this without support.
A single species is more likely to have consistent behavior, eating in one place and defecating in another, concentrating nutrients rather than dispersing them.
“A single species is more likely to have consistent behavior, eating in one place, defecating in another, and concentrating nutrients instead of dispersing them across gradients.”
The paper relies on this without support.
Multiple species have different eating and defecating patterns and are more likely to diffuse nutrients across gradients.
“By contrast, multiple species have different eating and defecating patterns and are more likely to diffuse nutrients across gradients.”
The paper relies on this without support.
Aquatic algae conduct most of the ocean's photosynthesis and have a much faster turnover time than land plants due to their often single-cellular nature.
“Aquatic algae, which conduct most of the ocean ' s photosynthesis, have a much faster turnover time than land plants due to their often single-cellular nature”
The authors state this as an assumption.
Twenty percent of guano arrives on land.
“assuming 20% of the guano arrives on land”
Claims other papers make about this one
Each is checked against this paper’s own text: does it say what the citing paper says it does?
The loss of large migrations of terrestrial vertebrates is associated with a loss in ecosystem functions they maintained.
“with an associated loss in ecosystem functions they maintained (1,41)”
From Annual mass drownings of the Serengeti wildebeest migration influence nutrient cycling and storage in the Mara River (2017) · cites this paper as [1]
The Discussion (p. 4) links the loss of terrestrial megafauna to a substantial reduction in animals’ capacity to distribute nutrients, an ecosystem function. The paper does not specifically establish that large migrations of terrestrial vertebrates were lost; it assesses extinctions and declines in large animals and the resulting loss of nutrient transport capacity. (LLM, conf=0.84)
Experimental — accuracy not yet measured
Self-check
Percentages, sums and reported differences re-computed from the paper’s own numbers.
Each contradiction between two passages listed here was confirmed by an independent check.
Africa's reported percentage does not match the table values
MinorTable 1 gives Africa a current capacity of 6.67e4 and a past capacity of 1.28e5. Dividing current by past gives 52.1%, not the reported 46%.
Reported 46 · recomputed 52.109375
“| % | 8% | 5% | 46%”
South America was not the highest-capacity continent in the table
MinorThe discussion says South America had the largest past nutrient distribution capacity, but Table 1 lists 1.53e5 for South America and a higher value, 2.77e5, for Eurasia.
“South America once had the largest nutrient distribution capacity”
“Land mammals 2.77e5 | Land mammals 2.39e5 | Land mammals 1.53e5”
The fish phosphorus flux is given diffusivity units
MinorThe results attach km2 per year to the historical 140-million phosphorus flux from anadromous fish. Table 1 gives that same quantity in kg P per year.
“anadromous fish may have moved at least an order of magnitude more P from ocean to land [140 million (71 -430 million) km 2 · y -1 ; SI Appendix , Table 2]”
“Past total, kg P · y - 1 | A. fish 1.4e8 kg · y - 1 Seabirds 6.3e6 ± 5e6”
The seabird total is assigned a per-area unit
MinorThe results describe the 6.3-million seabird flux in kg per square kilometer per year, whereas Table 1 presents the same figure as a total in kg P per year.
“We calculate a global flux of 6.3 million (1.5 -16 million; SI Appendix , Table 2) kg km -2 · y -1 of P from sea to land by the seabirds”
“Past total, kg P · y - 1 | A. fish 1.4e8 kg · y - 1 Seabirds 6.3e6 ± 5e6”
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Self-citationFishing down nutrients on coral reefs
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Self-citationIntegral functions of marine vertebrates in the ocean carbon cycle and climate change mitigation
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Self-citation
- Authors
- Doughty, C. E., Roman, J., Faurby, S., Wolf, A., Haque, A., Bakker, E. S., Malhi, Y., Dunning, J. B., & Svenning, J.-C.
- Affiliations
- University of Oxford (GB)
- University of Vermont (US)
- Aarhus University (DK)
- Princeton University (US)
- Netherlands Institute of Ecology (NL)
- Purdue University West Lafayette (US)
- ORCID
- Venue
- Proceedings of the National Academy of Sciences
- Publisher
- National Academy of Sciences
- Published
- 2015-10-26
- Volume
- 113
- Issue
- 4
- Pages
- 868-873
- ISSN
- 0027-8424 (ISSN-L)
- 1091-6490
- PMC
- PMC4743783
- OpenAlex
- W1826044145
- Semantic Scholar
- 73ec212cf94b82ed41724cc153e72b1e98c69f2b
- Cited by
- 470
- References
- 82
- Citations by year
- 2026: 27 · 2025: 46 · 2024: 50 · 2023: 63 · 2022: 46 · 2021: 49 · 2020: 63 · 2019: 47 · 2018: 26 · 2017: 32 · 2016: 17 · 2015: 3
- FWCI
- 23.57
- Type
- journal article
- Language
- en
- Open access
- bronze
- doi.org
- Locations
- Proceedings of the National Academy of Sciences (publishedVersion)
- PubMed Central (publishedVersion)
- PubMed (publishedVersion)
- KNAW Research Portal (The Royal Netherlands Academy of Arts and Sciences) (publishedVersion, other-oa)
- Data Archiving and Networked Services (DANS) (submittedVersion, other-oa)
- Funders
- Danmarks Grundforskningsfond
- Topics
- Physical Sciences › Environmental Science › Ecology › Marine animal studies overview
- Physical Sciences › Earth and Planetary Sciences › Oceanography › Marine and coastal ecosystems
- Physical Sciences › Environmental Science › Ecology › Animal Ecology and Behavior Studies
- Keywords
- nutrient cycling, megafaunal extinction, marine mammals, whales, seabirds, anadromous fish, phosphorus, nutrient transport, nutrient hotspots, large animals
- MeSH terms
- Animals, Behavior, Animal, Birds, Diffusion, Feeding Behavior, Fishes, Food, Food Supply, Homing Behavior, Mammals, Oceans and Seas, Population Density, Ecosystem, Body Size, History, Ancient, Extinction, Biological, Aquatic Organisms, Animal Distribution
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
Claims read by deepseek-v4.1-flash under definition version 3 (extract-v3) on 2026-10-04.
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