Deep-sea gigantism

Tendency for deep-sea species to be larger than their shallower-water relatives

Examination of a 9 m (30 ft) giant squid, the second largest cephalopod, that washed ashore in Norway in 1954

In zoology, deep-sea gigantism or abyssal gigantism is the tendency for species of deep-sea dwelling animals to be larger than their shallower-water relatives across a large taxonomic range. Proposed explanations for this type of gigantism include necessary adaptation to colder temperature, food scarcity, reduced predation pressure and increased dissolved oxygen concentrations in the deep sea. The harsh conditions and inhospitality of the underwater environment in general, as well as the inaccessibility of the abyssal zone for most human-made underwater vehicles, have hindered the study of this topic.

## Taxonomic range

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In marine crustaceans, the trend of increasing size with depth has been observed in mysids, euphausiids, decapods, isopods, ostracods and amphipods.[1][2] Non-arthropods in which deep-sea gigantism has been observed are cephalopods, cnidarians, and eels from the order Anguilliformes.[3][4]

> Other [animals] attain under them gigantic proportions. It is especially certain crustacea which exhibit this latter peculiarity, but not all crustacea, for the crayfish like forms in the deep sea are of ordinary size. I have already referred to a gigantic Pycnogonid [sea spider] dredged by us. Louis Agassiz dredged a gigantic Isopod 11 inches [28 centimetres] in length. We also dredged a gigantic Ostracod. – Henry Nottidge Moseley, 1880[5]

Notable organisms that exhibit deep-sea gigantism include the big red jellyfish,[6] Stygiomedusa jellyfish, the giant isopod,[5] giant ostracod,[5] the giant sea spider,[5] the giant amphipod, the Japanese spider crab, the giant oarfish, the deepwater stingray, the seven-arm octopus,[7] and a number of squid species: the colossal squid (up to 14 m in length),[8] the giant squid (up to 12 m),[8] Megalocranchia fisheri, robust clubhook squid, Dana octopus squid, cockatoo squid, giant warty squid, and the bigfin squids of the genus Magnapinna.

Deep-sea gigantism is not generally observed in the meiofauna (organisms that pass through a 1 mm (0.039 in) mesh), which actually exhibit the reverse trend of decreasing size with depth.[9]

## Explanations

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### Lower temperature

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In crustaceans, it has been proposed that the explanation for the increase in size with depth is similar to that for the increase in size with latitude (Bergmann's rule): both trends involve increasing size with decreasing temperature.[1] The trend with latitude has been observed in some of the same groups, both in comparisons of related species, as well as within widely distributed species.[1] Decreasing temperature is thought to result in increased cell size and increased life span (the latter also being associated with delayed sexual maturity[9]), both of which lead to an increase in maximum body size (continued growth throughout life is characteristic of crustaceans).[1] In Arctic and Antarctic seas where there is a reduced vertical temperature gradient, there is also a reduced trend towards increased body size with depth, arguing against hydrostatic pressure being an important parameter.[1]

Temperature does not appear to have a similar role in influencing the size of giant tube worms. Riftia pachyptila, which lives in hydrothermal vent communities at ambient temperatures of 2–30 °C,[10] reaches lengths of 2.7 m, comparable to those of Lamellibrachia luymesi, which lives in cold seeps. The former, however, has rapid growth rates and short life spans of about 2 years,[11] while the latter is slow growing and may live over 250 years.[12]

### Food scarcity

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Food scarcity at depths greater than 400 m is also thought to be a factor, since larger body size can improve ability to forage for widely scattered resources.[9] In organisms with planktonic eggs or larvae, another possible advantage is that larger offspring, with greater initial stored food reserves, can drift for greater distances.[9] As an example of adaptations to this situation, giant isopods gorge on food when available, distending their bodies to the point of compromising ability to locomote;[13] they can also survive 5 years without food in captivity.[14][15]

According to Kleiber's law,[16] the larger an animal gets, the more efficient its metabolism becomes; i.e., an animal's basal metabolic rate scales to roughly the ¾ power of its mass. Under conditions of limited food supply, this may provide additional benefit to large size.

### Reduced predation pressure

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An additional possible influence is reduced predation pressure in deeper waters.[17] A study of brachiopods found that predation was nearly an order of magnitude less frequent at the greatest depths than in shallow waters.[17]

### Increased dissolved oxygen

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Dissolved oxygen levels are also thought to play a role in deep-sea gigantism. A 1999 study of benthic amphipod crustaceans found that maximum potential organism size directly correlates with the increased levels of dissolved oxygen levels in deeper waters.[18] The solubility of dissolved oxygen in the oceans is known to lower in oxygen-poor intermediary depths (ranging 200-1000 meters) until the increasing pressure, decreasing salinity levels, and colder temperatures of deeper water can contribute increasing solubility once more.[18]

However, solubility does not necessarily equate to access as many areas of colder waters have exhibited lower levels of available oxygen. The proposed theory behind this trend is that deep-sea gigantism could be an adaptive trait to combat asphyxiation in frigid, dense ocean waters.[19] Larger organisms are able to intake more dissolved oxygen within the ocean, allowing for sufficient respiration. However, this increased absorption of oxygen runs the risk of toxicity poisoning where an organism can have oxygen levels that are so high that they become harmful and poisonous.[20]

## Impact of climate change

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Warmer global temperatures may have an effect on the deep sea as much as the shallower surface waters of the ocean, as evidence suggests that deep-sea ecosystems can be sensitive to shifts within the climate.[21] Despite appearing relatively unchanging, the dependence of life in the depths on food supply to drift down in the form of marine snow and water currents traveling throughout all depths of the ocean calls into concern the manner in which global climate change may affect these organisms and the biomes they live in.[21]

Following trends from the Paleocene-Eocene thermal maximum and related timescales, research suggests that current predictions of continuous greenhouse gas emissions and climate change will lead to higher ocean temperatures and a significant reduction in levels of dissolved oxygen in the deep sea.[22] Should global warming lead to a warmer ocean state, thermohaline circulation would no longer be able to maintain an oxygen-rich deep sea, which would eventually lead to deep water becoming higher in both temperature and salinity.[21]

Based upon current theories regarding the existence of deep-sea gigantism, we would expect to see the phenomenon diminish in response to these changes in the environment, as it may become unfavorable or even impossible for these organisms to sustain a larger body form.[22]

## Gallery

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* A giant isopod (Bathynomus giganteus) may reach up to 0.76 m (2 ft 6 in) in length.

* A Japanese spider crab whose outstretched legs measured 3.7 m (12 ft) across

* A robust clubhook squid, whose mantle reaches 2 m (6 ft 7 in) in length, caught off Alaska

* A 7 m (23 ft) king of herrings oarfish, washed up on the beach of a Navy SEAL training base in California

* A Colossendeis colossea sea spider, displayed at the Smithsonian

* A Stygiomedusa jellyfish, which can grow up to 10 m (33 ft) in length

* A deepwater stingray, which can reach up to 2.7 m × 1.5 m (8 ft 10 in × 4 ft 11 in) in size

## See also

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* Cephalopod size
* Dwarfing
* Island gigantism
* Insular dwarfism
* Largest organisms
* Megafauna

## References

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1. ^ a b c d e Timofeev, S. F. (2001). "Bergmann's Principle and Deep-Water Gigantism in Marine Crustaceans". Biology Bulletin of the Russian Academy of Sciences. 28 (6): 646–650\. Bibcode:2001BioBu..28..646T. doi:10.1023/A:1012336823275. S2CID 28016098.
2. ^ C., McClain; M., Rex (1 October 2001). "The relationship between dissolved oxygen concentration and maximum size in deep-sea turrid gastropods: an application of quantile regression". Marine Biology. 139 (4): 681–685\. Bibcode:2001MarBi.139..681C. doi:10.1007/s002270100617. ISSN 0025-3162. S2CID 83747571.
3. ^ Hanks, Micah. "Deep Sea Gigantism: Curious Cases of Mystery Giant Eels". MysteriousUniverse. Retrieved 5 May 2019.
4. ^ MacDonald, A.G. (1975). Physical Aspects of Deep Sea Biology. Cambridge University Press. pp. 17–19. ISBN 978-0-521-20397-5.
5. ^ a b c d McClain, Craig (14 January 2015). "Why isn't the Giant Isopod larger?". Deep Sea News. Retrieved 1 March 2018.
6. ^ Smithsonian Oceans. "Big Red Jellyfish". Smithsonian Oceans. Retrieved 5 May 2019.
7. ^ Hoving, H. J. T.; Haddock, S. H. D. (27 March 2017). "The giant deep-sea octopus Haliphron atlanticus forages on gelatinous fauna". Scientific Reports. 7: 44952. Bibcode:2017NatSR...744952H. doi:10.1038/srep44952. PMC 5366804. PMID 28344325.
8. ^ a b Anderton, Jim (22 February 2007). "Amazing specimen of world's largest squid in NZ". New Zealand Government. Archived from the original on 23 May 2010.
9. ^ a b c d Gad, G. (2005). "Giant Higgins-larvae with paedogenetic reproduction from the deep sea of the Angola Basin? Evidence for a new life cycle and for abyssal gigantism in Loricifera?". Organisms Diversity & Evolution. 5: 59–75\. Bibcode:2005ODivE...5...59G. doi:10.1016/j.ode.2004.10.005.
10. ^ Bright, M.; Lallier, F. H. (2010). The biology of vestimentiferan tubeworms (PDF). Oceanography and Marine Biology - an Annual Review. Vol. 48\. Taylor & Francis. pp. 213–266\. doi:10.1201/ebk1439821169. ISBN 978-1-4398-2116-9. Archived from the original (PDF) on 31 October 2013. Retrieved 30 October 2013.
11. ^ Lutz, R. A.; Shank, T. M.; Fornari, D. J.; Haymon, R. M.; Lilley, M. D.; Von Damm, K. L.; Desbruyeres, D. (1994). "Rapid growth at deep-sea vents". Nature. 371 (6499): 663. Bibcode:1994Natur.371..663L. doi:10.1038/371663a0. S2CID 4357672.
12. ^ MacDonald, Ian R. (2002). "Stability and Change in Gulf of Mexico Chemosynthetic Communities" (PDF). MMS. Archived from the original (PDF) on 1 February 2017. Retrieved 30 October 2013.
13. ^ Briones-Fourzán, Patricia; Lozano-Alvarez, Enrique (1991). "Aspects of the biology of the giant isopod Bathynomus giganteus A. Milne Edwards, 1879 (Flabellifera: Cirolanidae), off the Yucatan Peninsula". Journal of Crustacean Biology. 11 (3): 375–385\. Bibcode:1991JCBio..11..375B. doi:10.2307/1548464. JSTOR 1548464.
14. ^ Gallagher, Jack (26 February 2013). "Aquarium's deep-sea isopod hasn't eaten for over four years". The Japan Times. Retrieved 21 May 2013.
15. ^ "I Won't Eat, You Can't Make Me! (And They Couldn't)". NPR. 22 February 2014. Retrieved 23 February 2014.
16. ^ Kleiber, M. (1947). "Body Size and Metabolic Rate". Physiological Reviews. 27 (4): 511–541\. doi:10.1152/physrev.1947.27.4.511. PMID 20267758.
17. ^ a b Harper, E. M.; Peck, L. S. (2016). "Latitudinal and depth gradients in marine predation pressure". Global Ecology and Biogeography. 25 (6): 670–678\. Bibcode:2016GloEB..25..670H. doi:10.1111/geb.12444.
18. ^ a b Chapelle, Gauthier; Peck, Lloyd S. (1999). "Polar gigantism dictated by oxygen availability". Nature. 399 (6732): 114–115\. Bibcode:1999Natur.399..114C. doi:10.1038/20099. ISSN 0028-0836. S2CID 4308425.
19. ^ Verberk, Wilco C. E. P.; Atkinson, David (2013). "Why polar gigantism and Palaeozoic gigantism are not equivalent: effects of oxygen and temperature on the body size of ectotherms". Functional Ecology. 27 (6): 1275–1285\. Bibcode:2013FuEco..27.1275V. doi:10.1111/1365-2435.12152. hdl:2066/123399. ISSN 0269-8463. JSTOR 24033996. S2CID 5636563.
20. ^ Verberk, Wilco C. E. P.; Atkinson, David (2013). "Why polar gigantism and Palaeozoic gigantism are not equivalent: effects of oxygen and temperature on the body size of ectotherms". Functional Ecology. 27 (6): 1275–1285\. Bibcode:2013FuEco..27.1275V. doi:10.1111/1365-2435.12152. hdl:2066/123399. ISSN 0269-8463. JSTOR 24033996. S2CID 5636563.
21. ^ a b c Rogers, Alex David (2015). "Environmental Change in the Deep Ocean". Annual Review of Environment and Resources. 40: 1–38\. doi:10.1146/annurev-environ-102014-021415. ISSN 1543-5938.
22. ^ a b Yang, Jianing (2023). "A Prediction upon Deep-Sea Gigantism Characteristics of Crustaceans in Respond to Future Global Climate". In Li, Xiaolong; Yuan, Chunhui; Kent, John (eds.). Proceedings of the 6th International Conference on Economic Management and Green Development. Applied Economics and Policy Studies. Singapore: Springer Nature. pp. 147–159\. doi:10.1007/978-981-19-7826-5_14. ISBN 978-981-19-7826-5.

## External links

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* Science Daily: Midgets and giants in the deep sea

* v
* t
* e

Biological rules

Rules|

* Allen's rule Shorter appendages in colder climates
* Bateson's rule Extra limbs mirror their neighbours
* Bergmann's rule Larger bodies in colder climates
* Cope's rule Bodies get larger over time
* Deep-sea gigantism Larger bodies in deep-sea animals
* Dollo's law Loss of complex traits is irreversible
* Eichler's rule Parasites co-vary with their hosts
* Emery's rule Insect social parasites are often in same genus as their hosts
* Fahrenholz's rule Host and parasite phylogenies become congruent
* Foster's rule (Insular gigantism, Insular dwarfism) Small species get larger, large species smaller, after colonizing islands
* Gause's law Complete competitors cannot coexist
* Gloger's rule Lighter coloration in colder, drier climates
* Haldane's rule Hybrid sexes that are absent, rare, or sterile, are heterogamic
* Harrison's rule Parasites co-vary in size with their hosts
* Hamilton's rule Genes increase in frequency when relatedness of recipient to actor times benefit to recipient exceeds reproductive cost to actor
* Kleiber's law An animals metabolic rate decreases with its size
* Hennig's progression rule In cladistics, the most primitive species are found in earliest, central, part of group's area
* Jarman–Bell principle The correlation between the size of an animal and its diet quality; larger animals can consume lower quality diet
* Jordan's rule Inverse relationship between water temperature and no. of fin rays, vertebrae
* Lack's principle Birds lay only as many eggs as they can provide food for
* Rapoport's rule Latitudinal range increases with latitude
* Rensch's rule Sexual size dimorphism increases with size when males are larger, decreases with size when females are larger
* Rosa's rule Groups evolve from character variation in primitive species to a fixed character state in advanced ones
* Schmalhausen's law A population at limit of tolerance in one aspect is vulnerable to small differences in any other aspect
* Thayer's law The top of an animals coloration is darker than the bottom
* Thorson's rule No. of eggs of benthic marine invertebrates decreases with latitude
* Van Valen's law Probability of extinction of a group is constant over time
* von Baer's laws Embryos start from a common form and develop into increasingly specialised forms
* Williston's law Parts in an organism become reduced in number and specialized in function

Related|

* Countergradient variation Where genetics opposes environment as a factor
* Gigantothermy Large ectothermic animals more easily maintain constant body temperature

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