Deep-sea Gigantism
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In
zoology Zoology ( , ) is the scientific study of animals. Its studies include the anatomy, structure, embryology, Biological classification, classification, Ethology, habits, and distribution of all animals, both living and extinction, extinct, and ...
, 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 In biology, adaptation has three related meanings. Firstly, it is the dynamic evolutionary process of natural selection that fits organisms to their environment, enhancing their evolutionary fitness. Secondly, it is a state reached by the p ...
to colder temperature, food scarcity, reduced predation pressure and increased
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It can ...
concentrations in the
deep sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...
. The harsh conditions and inhospitality of the underwater environment in general, as well as the inaccessibility of the
abyssal zone The abyssal zone or abyssopelagic zone is a layer of the pelagic zone of the ocean. The word ''abyss'' comes from the Greek word (), meaning "bottomless". At depths of , this zone remains in perpetual darkness. It covers 83% of the total area ...
for most human-made
underwater vehicle An underwater vehicle is any member of the class of watercraft that is intended to operate in the underwater environment, as opposed to surface vessels that only stay afloat on top of water. This article lists the types of underwater vehicle, wit ...
s, have hindered the study of this topic.


Taxonomic range

In marine
crustacean Crustaceans (from Latin meaning: "those with shells" or "crusted ones") are invertebrate animals that constitute one group of arthropods that are traditionally a part of the subphylum Crustacea (), a large, diverse group of mainly aquatic arthrop ...
s, the trend of increasing size with depth has been observed in
mysid Mysida is an order of small, shrimp-like crustaceans in the malacostracan superorder Peracarida. Their common name opossum shrimps stems from the presence of a brood pouch or "marsupium" in females. The fact that the larvae are reared in thi ...
s, euphausiids,
decapod The Decapoda or decapods, from Ancient Greek δεκάς (''dekás''), meaning "ten", and πούς (''poús''), meaning "foot", is a large order of crustaceans within the class Malacostraca, and includes crabs, lobsters, crayfish, shrimp, and p ...
s,
isopod Isopoda is an Order (biology), order of crustaceans. Members of this group are called isopods and include both Aquatic animal, aquatic species and Terrestrial animal, terrestrial species such as woodlice. All have rigid, segmented exoskeletons ...
s,
ostracod Ostracods, or ostracodes, are a Class (biology), class of the crustacean, Crustacea (class Ostracoda), sometimes known as seed shrimp. Some 33,000 species (only 13,000 of which are extant taxon, extant) have been identified,Brandão, S.N.; Antoni ...
s and amphipods. Non-arthropods in which deep-sea gigantism has been observed are
cephalopod A cephalopod is any member of the molluscan Taxonomic rank, class Cephalopoda (Greek language, Greek plural , ; "head-feet") such as a squid, octopus, cuttlefish, or nautilus. These exclusively marine animals are characterized by bilateral symm ...
s,
cnidaria Cnidaria ( ) is a phylum under kingdom Animalia containing over 11,000 species of aquatic invertebrates found both in fresh water, freshwater and marine environments (predominantly the latter), including jellyfish, hydroid (zoology), hydroids, ...
ns, and eels from the order
Anguilliformes Eels are ray-finned fish belonging to the order (biology), order Anguilliformes (), which consists of eight suborders, 20 Family (biology), families, 164 genus, genera, and about 1000 species. Eels undergo considerable development from the earl ...
. Notable organisms that exhibit deep-sea gigantism include the big red jellyfish, '' Stygiomedusa''
jellyfish Jellyfish, also known as sea jellies or simply jellies, are the #Life cycle, medusa-phase of certain gelatinous members of the subphylum Medusozoa, which is a major part of the phylum Cnidaria. Jellyfish are mainly free-swimming marine animal ...
, the giant isopod, giant ostracod, the giant sea spider, the giant amphipod, the Japanese spider crab, the giant oarfish, the deepwater stingray, the seven-arm octopus, and a number of
squid A squid (: squid) is a mollusc with an elongated soft body, large eyes, eight cephalopod limb, arms, and two tentacles in the orders Myopsida, Oegopsida, and Bathyteuthida (though many other molluscs within the broader Neocoleoidea are also ...
species: the colossal squid (up to 14 m in length), the giant squid (up to 12 m), '' 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 mesh), which actually exhibit the reverse trend of decreasing size with depth.


Explanations


Lower temperature

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 In geography, latitude is a geographic coordinate system, geographic coordinate that specifies the north-south position of a point on the surface of the Earth or another celestial body. Latitude is given as an angle that ranges from −90° at t ...
( Bergmann's rule): both trends involve increasing size with decreasing temperature. 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. Decreasing temperature is thought to result in increased cell size and increased life span (the latter also being associated with delayed sexual maturity), both of which lead to an increase in maximum body size (continued growth throughout life is characteristic of crustaceans). In
Arctic The Arctic (; . ) is the polar regions of Earth, polar region of Earth that surrounds the North Pole, lying within the Arctic Circle. The Arctic region, from the IERS Reference Meridian travelling east, consists of parts of northern Norway ( ...
and
Antarctic The Antarctic (, ; commonly ) is the polar regions of Earth, polar region of Earth that surrounds the South Pole, lying within the Antarctic Circle. It is antipodes, diametrically opposite of the Arctic region around the North Pole. The Antar ...
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 Hydrostatics is the branch of fluid mechanics that studies fluids at hydrostatic equilibrium and "the pressure in a fluid or exerted by a fluid on an immersed body". The word "hydrostatics" is sometimes used to refer specifically to water and o ...
being an important parameter. Temperature does not appear to have a similar role in influencing the size of giant tube worms. '' Riftia pachyptila'', which lives in
hydrothermal vent Hydrothermal vents are fissures on the seabed from which geothermally heated water discharges. They are commonly found near volcanically active places, areas where tectonic plates are moving apart at mid-ocean ridges, ocean basins, and hot ...
communities at ambient temperatures of 2–30 °C, reaches lengths of 2.7 m, comparable to those of '' Lamellibrachia luymesi'', which lives in
cold seep A cold seep (sometimes called a cold vent) is an area of the ocean floor where seepage of fluids rich in hydrogen sulfide, methane, and other hydrocarbons occurs, often in the form of a brine pool. ''Cold'' does not mean that the temperature ...
s. The former, however, has rapid growth rates and short life spans of about 2 years, while the latter is slow growing and may live over 250 years.


Food scarcity

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. In organisms with
plankton Plankton are the diverse collection of organisms that drift in Hydrosphere, water (or atmosphere, air) but are unable to actively propel themselves against ocean current, currents (or wind). The individual organisms constituting plankton are ca ...
ic eggs or larvae, another possible advantage is that larger offspring, with greater initial stored food reserves, can drift for greater distances. 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; they can also survive 5 years without food in captivity. According to Kleiber's law, the larger an animal gets, the more efficient its metabolism becomes; i.e., an animal's
basal metabolic rate Basal metabolic rate (BMR) is the rate of energy expenditure per unit time by endothermic animals at rest.. In other words it is the energy required by body organs to perform normal It is reported in energy units per unit time ranging from watt ( ...
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

An additional possible influence is reduced predation pressure in deeper waters. A study of
brachiopod Brachiopods (), phylum (biology), phylum Brachiopoda, are a phylum of animals that have hard "valves" (shells) on the upper and lower surfaces, unlike the left and right arrangement in bivalve molluscs. Brachiopod valves are hinged at the rear e ...
s found that predation was nearly an order of magnitude less frequent at the greatest depths than in shallow waters.


Increased dissolved oxygen

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. 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. 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. 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.


Impact of Climate Change

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. 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. 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. Should global warming lead to a warmer ocean state,
thermohaline circulation Thermohaline circulation (THC) is a part of the large-scale Ocean current, ocean circulation driven by global density gradients formed by surface heat and freshwater fluxes. The name ''thermohaline'' is derived from ''wikt:thermo-, thermo-'', r ...
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. 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.


Gallery

File:Giant isopod.jpg, A giant isopod (''Bathynomus giganteus'') may reach up to in length. File:Japanese spider crab.jpg, A Japanese spider crab whose outstretched legs measured across File:Moroteuthis robusta bronco.jpg, A robust clubhook squid, whose mantle reaches in length, caught off
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
File:Giant Oarfish.jpg, A king of herrings oarfish, washed up on the beach of a Navy SEAL training base in
California California () is a U.S. state, state in the Western United States that lies on the West Coast of the United States, Pacific Coast. It borders Oregon to the north, Nevada and Arizona to the east, and shares Mexico–United States border, an ...
File:Colossendeis colossea Smithsonian Natural History Museum.jpg, A '' Colossendeis colossea'' sea spider, displayed at the Smithsonian File:Stygiomedusa Gigantea ov.jpg, A '' Stygiomedusa'' jellyfish, which can grow up to in length File:Plesiobatis daviesi cochin.jpg, A deepwater stingray, which can reach up to in size


See also

*
Cephalopod size Cephalopods, which include squids and octopuses, vary enormously in size. The smallest are only about long and weigh less than at maturity, while the giant squid can exceed in length and the colossal squid weighs close to half a tonne (), makin ...
* Dwarfing * Island gigantism *
Insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
* Largest organisms *
Megafauna In zoology, megafauna (from Ancient Greek, Greek μέγας ''megas'' "large" and Neo-Latin ''fauna'' "animal life") are large animals. The precise definition of the term varies widely, though a common threshold is approximately , this lower en ...


References


External links


Science Daily: Midgets and giants in the deep sea
{{DEFAULTSORT:Deep-Sea Gigantism Animal size Evolutionary biology concepts Marine organisms Ecogeographic rules