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Lanternfishes (or myctophids, from the
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
μυκτήρ ''myktḗr'', "nose" and ''ophis'', "serpent") are small mesopelagic fish of the large
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Ideal ...
Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represented by 246
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
in 33
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencla ...
, and are found in oceans worldwide.
Lantern A lantern is an often portable source of lighting, typically featuring a protective enclosure for the light sourcehistorically usually a candle or a wick in oil, and often a battery-powered light in modern timesto make it easier to carry and h ...
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of liv ...
es are aptly named after their conspicuous use of
bioluminescence Bioluminescence is the production and emission of light by living organisms. It is a form of chemiluminescence. Bioluminescence occurs widely in marine vertebrates and invertebrates, as well as in some fungi, microorganisms including some ...
. Their sister family, the Neoscopelidae, are much fewer in number but superficially very similar; at least one neoscopelid shares the
common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) is a name that is based on the normal language of everyday life; and is often contrast ...
"lanternfish": the large-scaled lantern fish, '' Neoscopelus macrolepidotus''. Lanternfish are among the most widely distributed, diverse and populous
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, wi ...
s, with some estimates suggesting that they may have a total global
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
of 1.8 to 16
gigatonne The tonne ( or ; symbol: t) is a unit of mass equal to 1000 kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton (United States c ...
s, accounting for up to 65% of all deep-sea fish biomass. Commercial fisheries for them exist off
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring coun ...
, in the sub-Antarctic, and in the
Gulf of Oman The Gulf of Oman or Sea of Oman ( ar, خليج عمان ''khalīj ʿumān''; fa, دریای عمان ''daryâ-ye omân''), also known as Gulf of Makran or Sea of Makran ( ar, خلیج مکران ''khalīj makrān''; fa, دریای مکرا� ...
.


Description

Lanternfish typically have a slender, compressed body covered in small, silvery deciduous
cycloid scale A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as ...
s (
ctenoid A fish scale is a small rigid plate that grows out of the skin of a fish. The skin of most jawed fishes is covered with these protective scales, which can also provide effective camouflage through the use of reflection and colouration, as ...
in four species), a large bluntly rounded head, large elliptical to round lateral eyes (dorsolateral in '' Protomyctophum'' species), and a large terminal mouth with jaws closely set with rows of small teeth. The fins are generally small, with a single high
dorsal fin A dorsal fin is a fin located on the back of most marine and freshwater vertebrates within various taxa of the animal kingdom. Many species of animals possessing dorsal fins are not particularly closely related to each other, though through co ...
, a forked
caudal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as s ...
, and an
adipose fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
. The
anal fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
is supported by a
cartilaginous Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck and ...
plate at its base, and originates under, or slightly behind, the rear part of the dorsal fin. The
pectoral fin Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
s, usually with eight rays, may be large and well-developed to small and degenerate, or completely absent in a few species. In some species, such as those of the genus '' Lampanyctus'', the pectorals are greatly elongated. Most lanternfish have a
gas bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their current water depth wi ...
, but it degenerates or fills with
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...
s during the maturation of a few species. The
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelial ...
is uninterrupted. In all but one species, ''Taaningichthys paurolychnus'', a number of
photophore A photophore is a glandular organ that appears as luminous spots on various marine animals, including fish and cephalopods. The organ can be simple, or as complex as the human eye; equipped with lenses, shutters, color filters and reflectors, ...
s (light-producing organs) are present; these are paired and concentrated in ventrolateral rows on the body and head. Some may also possess specialised photophores on the
caudal peduncle Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as se ...
, in proximity to the eyes (e.g., the "headlights" of '' Diaphus'' species), and luminous patches at the base of the fins. The photophores emit a weak blue, green, or yellow light, and are known to be arranged in species-specific patterns. In some species, the pattern varies between males and females. This is true for the luminous caudal patches, with the males' being typically above the tail and the females' being below the tail. Lanternfish are generally small fish, ranging from about in length, with most being under . Shallow-living species are an
iridescent Iridescence (also known as goniochromism) is the phenomenon of certain surfaces that appear to gradually change color as the angle of view or the angle of illumination changes. Examples of iridescence include soap bubbles, feathers, butterfly ...
blue to green or silver, while deeper-living species are dark brown to black.


Ecology

Lanternfish are well known for their
diel vertical migration Diel vertical migration (DVM), also known as diurnal vertical migration, is a pattern of movement used by some organisms, such as copepods, living in the ocean and in lakes. The word ''diel'' comes from the Latin ''dies'' day, and means a 24- ...
s: during daylight hours, most species remain within the gloomy bathypelagic zone, between deep, but towards sundown, the fish begin to rise into the epipelagic zone, between deep. The lanternfish are thought to do this to avoid predation, and because they are following the diel vertical migrations of
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
, upon which they feed. After a night spent feeding in the surface layers of the water column, the lanternfish begin to descend back into the lightless depths and are gone by daybreak. By releasing fecal pellets at depth, Laternfish make the carbon capture process called
biological pump The biological pump (or ocean carbon biological pump or marine biological carbon pump) is the ocean's biologically driven sequestration of carbon from the atmosphere and land runoff to the ocean interior and seafloor sediments.Sigman DM & GH ...
more efficient. Most species remain near the coast,
schooling A school is an educational institution designed to provide learning spaces and learning environments for the teaching of students under the direction of teachers. Most countries have systems of formal education, which is sometimes compulsor ...
over the
continental slope A continental margin is the outer edge of continental crust abutting oceanic crust under coastal waters. It is one of the three major zones of the ocean floor, the other two being deep-ocean basins and mid-ocean ridges. The continental margin ...
. Different species are known to segregate themselves by depth, forming dense, discrete conspecific layers, probably to avoid competition between different species. Due to their gas bladders, these layers are visible on sonar scans and give the impression of a "false bottom"; this is the so-called
deep scattering layer The deep scattering layer, sometimes referred to as the sound scattering layer, is a layer in the ocean consisting of a variety of marine animals. It was discovered through the use of sonar, as ships found a layer that scattered the sound and w ...
that so perplexed early oceanographers (see below). Great variability in migration patterns occurs within the family. Some deeper-living species may not migrate at all, while others may do so only sporadically. Migration patterns may also depend on life stage, sex,
latitude In geography, latitude is a 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 the south pole to 90° at the north ...
, and season. The arrangements of lanternfish photophores are different for each species, so their bioluminescence is thought to play a role in
communication Communication (from la, communicare, meaning "to share" or "to be in relation with") is usually defined as the transmission of information. The term may also refer to the message communicated through such transmissions or the field of inqu ...
, specifically in shoaling and
courtship Courtship is the period wherein some couples get to know each other prior to a possible marriage. Courtship traditionally may begin after a betrothal and may conclude with the celebration of marriage. A courtship may be an informal and private ...
behaviour. The concentration of the photophores on the flanks of the fish also indicate the light's use as
camouflage Camouflage is the use of any combination of materials, coloration, or illumination for concealment, either by making animals or objects hard to see, or by disguising them as something else. Examples include the leopard's spotted coat, the b ...
; in a strategy termed counterillumination, the lanternfish regulate the brightness of the bluish light emitted by their photophores to match the ambient light level above, effectively masking the lanternfishes' silhouette when viewed from below. A major source of food for many marine animals, lanternfish are an important link in the
food chain A food chain is a linear network of links in a food web starting from producer organisms (such as grass or algae which produce their own food via photosynthesis) and ending at an apex predator species (like grizzly bears or killer whales), det ...
of many local
ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...
s, being heavily preyed upon by whales and
dolphin A dolphin is an aquatic mammal within the infraorder Cetacea. Dolphin species belong to the families Delphinidae (the oceanic dolphins), Platanistidae (the Indian river dolphins), Iniidae (the New World river dolphins), Pontoporiidae ( ...
s, large
pelagic fish Pelagic fish live in the pelagic zone of ocean or lake waters—being neither close to the bottom nor near the shore—in contrast with demersal fish that do live on or near the bottom, and reef fish that are associated with coral reef ...
such as
salmon Salmon () is the common name for several commercially important species of euryhaline ray-finned fish from the family Salmonidae, which are native to tributaries of the North Atlantic (genus ''Salmo'') and North Pacific (genus '' Oncorhync ...
,
tuna A tuna is a saltwater fish that belongs to the tribe Thunnini, a subgrouping of the Scombridae ( mackerel) family. The Thunnini comprise 15 species across five genera, the sizes of which vary greatly, ranging from the bullet tuna (max len ...
and
shark Sharks are a group of elasmobranch fish characterized by a cartilaginous skeleton, five to seven gill slits on the sides of the head, and pectoral fins that are not fused to the head. Modern sharks are classified within the clade Selachimo ...
s,
grenadier A grenadier ( , ; derived from the word ''grenade'') was originally a specialist soldier who threw hand grenades in battle. The distinct combat function of the grenadier was established in the mid-17th century, when grenadiers were recruited from ...
s and other deep-sea fish (including other lanternfish),
pinniped Pinnipeds (pronounced ), commonly known as seals, are a widely distributed and diverse clade of carnivorous, fin-footed, semiaquatic, mostly marine mammals. They comprise the extant families Odobenidae (whose only living member is the walr ...
s,
sea bird Seabirds (also known as marine birds) are birds that are adapted to life within the marine environment. While seabirds vary greatly in lifestyle, behaviour and physiology, they often exhibit striking convergent evolution, as the same enviro ...
s, notably
penguin Penguins ( order Sphenisciformes , family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain t ...
s, and large
squid True squid are molluscs with an elongated soft body, large eyes, eight arms, and two tentacles in the superorder Decapodiformes, though many other molluscs within the broader Neocoleoidea are also called squid despite not strictly fitting ...
such as the jumbo squid, ''Dosidicus gigas''. Lanternfish themselves have been found to feed on bits of plastic debris accumulating in the oceans. At least one lanternfish was found with over 80 pieces of plastic chips in its gut, according to scientists monitoring ocean plastic in the Pacific Ocean's eastern garbage patch.


Deep scattering layer

Sonar operators, using the newly developed sonar technology during World War II, were puzzled by what appeared to be a false sea floor 300–500 metres deep at day, and less deep at night. This turned out to be due to millions of marine organisms, most particularly small
mesopelagic The mesopelagic zone ( Greek μέσον, middle), also known as the middle pelagic or twilight zone, is the part of the pelagic zone that lies between the photic epipelagic and the aphotic bathypelagic zones. It is defined by light, and begins a ...
fish, with swimbladders that reflected the sonar. These organisms migrate up into shallower water at dusk to feed on
plankton Plankton are the diverse collection of organisms found in water (or air) that are unable to propel themselves against a current (or wind). The individual organisms constituting plankton are called plankters. In the ocean, they provide a cruci ...
. The layer is deeper when the moon is out, and can become shallower when clouds pass over the moon.Ryan
"Deep-sea creatures: The mesopelagic zone"
''Te Ara - the Encyclopedia of New Zealand''. Updated 21 September 2007.
Sampling via deep
trawling Trawling is a method of fishing that involves pulling a fishing net through the water behind one or more boats. The net used for trawling is called a trawl. This principle requires netting bags which are towed through water to catch different speci ...
indicates that lanternfish account for as much as 65% of all deep sea fish
biomass Biomass is plant-based material used as a fuel for heat or electricity production. It can be in the form of wood, wood residues, energy crops, agricultural residues, and waste from industry, farms, and households. Some people use the terms bi ...
. Indeed, lanternfish are among the most widely distributed, populous, and diverse of all
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, wi ...
s, playing an important
ecological Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overlaps w ...
role as prey for larger organisms. The estimated global biomass of lanternfish is 550–660 million
tonne The tonne ( or ; symbol: t) is a unit of mass equal to 1000  kilograms. It is a non-SI unit accepted for use with SI. It is also referred to as a metric ton to distinguish it from the non-metric units of the short ton (United States ...
s, several times the entire world fisheries catch. Lanternfish also account for much of the biomass responsible for the
deep scattering layer The deep scattering layer, sometimes referred to as the sound scattering layer, is a layer in the ocean consisting of a variety of marine animals. It was discovered through the use of sonar, as ships found a layer that scattered the sound and w ...
of the world's oceans. Sonar reflects off the millions of lanternfish
swim bladder The swim bladder, gas bladder, fish maw, or air bladder is an internal gas-filled organ that contributes to the ability of many bony fish (but not cartilaginous fish) to control their buoyancy, and thus to stay at their current water depth wi ...
s, giving the appearance of a false bottom.


Rise to dominance

Lanternfish currently represent one of the dominant groups of mesopelagic fishes in terms of abundance, biomass, and diversity. Their
otolith An otolith ( grc-gre, ὠτο-, ' ear + , ', a stone), also called statoconium or otoconium or statolith, is a calcium carbonate structure in the saccule or utricle of the inner ear, specifically in the vestibular system of vertebrates. The sa ...
record dominates
pelagic sediment Pelagic sediment or pelagite is a fine-grained sediment that accumulates as the result of the settling of particles to the floor of the open ocean, far from land. These particles consist primarily of either the microscopic, calcareous or siliceou ...
s below 200 m in dredges, especially during the entire
Neogene The Neogene ( ), informally Upper Tertiary or Late Tertiary, is a geologic period and system that spans 20.45 million years from the end of the Paleogene Period million years ago ( Mya) to the beginning of the present Quaternary Period Mya. ...
. The diversity and rise to dominance of lanternfish can be examined by analysing these otolith records. The earliest unambiguous fossil lanternfish are known based on otoliths from the
late Paleocene The Thanetian is, in the ICS Geologic timescale, the latest age or uppermost stratigraphic stage of the Paleocene Epoch or Series. It spans the time between . The Thanetian is preceded by the Selandian Age and followed by the Ypresian Age (p ...
and
early Eocene In the geologic timescale the Ypresian is the oldest age or lowest stratigraphic stage of the Eocene. It spans the time between , is preceded by the Thanetian Age (part of the Paleocene) and is followed by the Eocene Lutetian Age. The Ypresia ...
. During their early evolutionary history, lanternfish were likely not adapted to a high oceanic lifestyle but occurred over shelf and upper-slope regions, where they were locally abundant during the
middle Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', "daw ...
. A distinct upscaling in otolith size is observed in the
early Oligocene The Rupelian is, in the geologic timescale, the older of two ages or the lower of two stages of the Oligocene Epoch/Series. It spans the time between . It is preceded by the Priabonian Stage (part of the Eocene) and is followed by the Chat ...
, which also marks their earliest occurrence in bathyal sediments. This transition is interpreted to be related to the change from a halothermal deep-ocean circulation to a
thermohaline Thermohaline circulation (THC) is a part of the large-scale ocean circulation that is driven by global density gradients created by surface heat and freshwater fluxes. The adjective ''thermohaline'' derives from '' thermo-'' referring to tempe ...
regime and the associated cooling of the deep ocean and rearrangement of nutrient and
silica Silicon dioxide, also known as silica, is an oxide of silicon with the chemical formula , most commonly found in nature as quartz and in various living organisms. In many parts of the world, silica is the major constituent of sand. Silica is ...
supply. The size of early Oligocene lanternfish is remarkably congruent with
diatom A diatom ( Neo-Latin ''diatoma''), "a cutting through, a severance", from el, διάτομος, diátomos, "cut in half, divided equally" from el, διατέμνω, diatémno, "to cut in twain". is any member of a large group comprising se ...
abundance, the main food resource for the
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
and thus for lanternfish and whales. The warmer
late Oligocene The Chattian is, in the geologic timescale, the younger of two ages or upper of two stages of the Oligocene Epoch/Series. It spans the time between . The Chattian is preceded by the Rupelian and is followed by the Aquitanian (the lowest stag ...
to early
middle Miocene The Middle Miocene is a sub-epoch of the Miocene Epoch made up of two stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0.05 Ma to 11.608 ± 0.005 Ma (million ...
period was characterised by an increase in the disparity of lanternfish but with a reduction in their otolith sizes. A second and persisting
secular Secularity, also the secular or secularness (from Latin ''saeculum'', "worldly" or "of a generation"), is the state of being unrelated or neutral in regards to religion. Anything that does not have an explicit reference to religion, either negativ ...
pulse in lanternfish diversity (particularly within the genus '' Diaphus'') and increase in size begins with the "biogenic bloom" during the
late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The e ...
, paralleled with diatom abundance and
gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...
in
baleen whale Baleen whales (systematic name Mysticeti), also known as whalebone whales, are a parvorder of carnivorous marine mammals of the infraorder Cetacea ( whales, dolphins and porpoises) which use keratinaceous baleen plates (or "whalebone") in the ...
s.


Genera

'' Benthosema''
'' Bolinichthys''
'' Centrobranchus''
'' Ceratoscopelus''
'' Diaphus''
'' Diogenichthys''
'' Electrona''
'' Gonichthys''
'' Gymnoscopelus''
'' Hintonia''
'' Hygophum''
'' Idiolychnus''
'' Krefftichthys''
'' Lampadena''
'' Lampanyctodes''
'' Lampanyctus''
'' Lampichthys''
'' Lepidophanes''
'' Lobianchia''
'' Loweina''
'' Metelectrona''
'' Myctophum''
'' Nannobrachium''
'' Notolychnus''
'' Notoscopelus''
'' Parvilux''
'' Protomyctophum''
'' Scopelopsis''
'' Stenobrachius''
'' Symbolophorus''
'' Taaningichthys''
'' Tarletonbeania''
'' Triphoturus''


References


Further references

* *
Order Myctophiformes: Blackchins and Lanternfishes
. Bethune-Cookman College, Moser, G. H., Watson, W. Retrieved December 13, 2004. (Retrieved from web archive July 10, 2006) *

. Iziko Museums of Cape Town. Hulley, P. A. Retrieved December 13, 2004. {{Authority control Ray-finned fish families Deep sea fish Bioluminescent fish Extant Cretaceous first appearances Taxa named by Theodore Gill