Stoplight loosejaw | |
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Illustration of a stoplight loosejaw | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Stomiiformes |
Family: | Stomiidae |
Subfamily: | Malacosteinae |
Genus: | Malacosteus Ayres, 1848 |
The stoplight loosejaws are small, deep-sea dragonfishes of the genus Malacosteus, classified either within the subfamily Malacosteinae of the family Stomiidae, or in the separate family Malacosteidae. They are found worldwide, outside of the Arctic and Subantarctic, in the mesopelagic zone below a depth of 500 meters (1,600 feet). This genus once contained three nominal species: M. niger (the type), M. choristodactylus, and M. danae, with the validity of the latter two species being challenged by different authors at various times. In 2007, Kenaley examined over 450 stoplight loosejaw specimens and revised the genus to contain two species, M. niger and the new M. australis. [1]
Malacosteus and the related genera Aristostomias , Chirostomias and Pachystomias are the only fishes that produce red bioluminescence. As most of their prey organisms are not capable of perceiving light at those wavelengths, this allows Malacosteus to hunt with an essentially invisible beam of light. Furthermore, Malacosteus is unique amongst animals in using a chlorophyll derivative to perceive red light. [1] The name Malacosteus is derived from the Greek malakos meaning "soft" and osteon meaning "bone". [2] Another common name for these fishes is "rat-trap fish", from the unusual open structure of their jaws. [3]
There are currently two recognized species in this genus: [4]
These fishes have a wide distribution in all oceans: M. niger is found between 66° N and 33° S, except for the Mediterranean Sea, while M. australis is found in the southern transition zone between 25° and 45° S, where it is bound by the Antarctic Circumpolar Current. M. niger appears to be replaced by M. australis south of 30° S, while M. australis does not occur north of that latitude outside of the Indian Ocean and the Indo-Australian Archipelago. Both species are usually found below a depth of 500 meters (1,600 feet) in midwater. They are the only known stomiids that do not seem to conduct significant diel vertical migrations. [1]
Malacosteus has an elongated body with short, blunt snouts and large eyes that face forward, granting binocular vision. Unlike other stomiids, it has a single round nostril on each side in front of the eye. Relative to its size, Malacosteus has one of the widest gapes of any fish, with a lower jaw measuring one-quarter of the fish's length. The lower jaw has no ethmoid membrane (floor) and is attached only by the hinge and a modified tongue bone. There are several large, fang-like teeth in the front of the jaws, followed by many small barbed teeth. There are several groups of pharyngeal teeth that serve to direct food down the esophagus. [1] [5]
The pectoral and pelvic fins are moderately long, containing 3–4 and 6 fin rays respectively. The dorsal and anal fins are placed far back on the body and contain 18–20 and 19–22 rays respectively. The caudal fin is small, with the lower lobe larger than the upper. There are three bioluminescent photophores near the eyes: beneath the eye is a large, teardrop-shaped suborbital photophore that emits red light. Behind it is an ovoid postorbital photophore that emits green light; this photophore is larger in males than females. These red and green photophores are evocative of traffic lights, hence the fish's common name. The third is tiny and round, located between the eye and the large red photophore. Several rows and clusters of blue photophores are present on the sides and belly. In addition, there are small photophores and accessory areas of white luminous tissue scattered over the head and body. The skin is thin and scaleless; the coloration is black. [1]
As long wavelengths of light (i.e. red) do not reach the deep sea from the surface, many deep-sea organisms are insensitive to red wavelengths, and so to these creatures, red-colored objects appear black. The red photophore of Malacosteus thus allows it to illuminate prey without being detected. These fishes exhibit a number of adaptations for feeding on large prey. The "open" structure of its jaws allows the fish to swing its entire head forward to grab prey from afar in addition to reducing water resistance, allowing them to be snapped shut more quickly, while large recurved teeth and powerful jaw muscles assure a secure hold. The connection between the head and the body is reduced, with unossified vertebrae, allowing the cranium to be tilted back and the jaws thrust forward for a wider gape. Finally, the gills are exposed to the outside, allowing the fish to continue respiring while slowly swallowing large prey. [6]
However, contrary to its apparent morphological specialization, the diet of Malacosteus consists primarily of zooplankton, chiefly large calanoid copepods, with smaller numbers of krill, shrimps, and fishes. It is yet unclear how Malacosteus captures such small planktonic prey given the open structure of its mouth. [5] The unexpected diet of Malacosteus is theorized to be a result of the small volumes that it searches for food, in which large prey items are rare. The rapid attenuation of red light in sea water gives Malacosteus a shorter visual range than species that use blue light, and it does not migrate vertically into more productive waters like other stomiids. Therefore, its strategy may be one of "snacking" on copepods, which are three orders of magnitude more abundant than fishes at its native depths, in between larger meals. [5]
The other factor believed to be partly responsible for Malacosteus' diet is its unique visual system, which uses a derivative of chlorophyll as a photosensitizer that absorbs long-wave light (around 700 nm) and then indirectly stimulates the fish's two visual pigments, which have maximum absorbances at only 520 and 540 nm. No vertebrates are known to synthesize chlorophyll derivatives, and Malacosteus is believed to obtain these derivatives from the copepods it consumes. [7] The red photophore of Malacosteus consists of a pigmented sac with a reflective inner lining and an internal mass of gland cells. Inside the gland cells, blue-green light is produced via the same chemical reaction found in other stomiids, which is then absorbed by a protein that fluoresces in a broad red band. This light is then reflected out through the photophore aperture, where it passes through a brown filter, yielding a far-red light with a maximum absorbance at 708 nm (almost infrared). In live fish, the suborbital and postorbital photophores both flash vigorously, the suborbital at a slower rate. [8] [9]
Deep-sea fish are fish that live in the darkness below the sunlit surface waters, that is below the epipelagic or photic zone of the sea. The lanternfish is, by far, the most common deep-sea fish. Other deep-sea fishes include the flashlight fish, cookiecutter shark, bristlemouths, anglerfish, viperfish, and some species of eelpout.
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 bioluminescent bacteria, and terrestrial arthropods such as fireflies. In some animals, the light is bacteriogenic, produced by symbiotic bacteria such as those from the genus Vibrio; in others, it is autogenic, produced by the animals themselves.
Stomiiformes is an order of deep-sea ray-finned fishes of very diverse morphology. It includes, for example, dragonfishes, lightfishes, loosejaws, marine hatchetfishes and viperfishes. The order contains 4 families with more than 50 genera and at least 410 species. As usual for deep-sea fishes, there are few common names for species of the order, but the Stomiiformes as a whole are often called dragonfishes and allies or simply stomiiforms.
Cetomimidae is a family of small, deep-sea cetomimiform fish. They are among the most deep-living fish known, with some species recorded at depths in excess of 3,500 m (11,500 ft). Females are known as flabby whalefishes, Males are known as bignose fishes, while juveniles are known as tapetails and were formerly thought to be in a separate family, dubbed Mirapinnidae. Adults exhibit extreme sexual dimorphism, and the adult males were once thought to be exemplars of still another family, Megalomycteridae.
Lanternfish are small mesopelagic fish of the large family Myctophidae. One of two families in the order Myctophiformes, the Myctophidae are represented by 246 species in 33 genera, and are found in oceans worldwide. Lanternfishes are aptly named after their conspicuous use of bioluminescence. Their sister family, the Neoscopelidae, are much fewer in number but superficially very similar; at least one neoscopelid shares the common name "lanternfish": the large-scaled lantern fish, Neoscopelus macrolepidotus.
The Pacific viperfish, Chauliodus macouni, is a predatory deep-sea fish found in the North Pacific. It is reported as being either mesopelagic or bathypelagic, with diel vertical migration to shallower waters. The Pacific viperfish is one of the nine different species that belong to the genus Chauliodus, the viperfish. The Pacific viperfish tend to be the largest of the species, typically reaching lengths of up to 1 foot and are considered an example of deep-sea gigantism. The length-weight relationship of the pacific viperfish varies with sex with females tending to be longer and heavier than males.
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 live on or near the bottom, and reef fish that are associated with coral reefs.
A viperfish is any species of marine fish in the genus Chauliodus. Viperfishes are mostly found in the mesopelagic zone and are characterized by long, needle-like teeth and hinged lower jaws. A typical viperfish grows to lengths of 30 cm (12 in). Viperfishes undergo diel vertical migration and are found all around the world in tropical and temperate oceans. Viperfishes are capable of bioluminescence and possess photophores along the ventral side of their body, likely used to camouflage them by blending in with the less than 1% of light that reaches to below 200 meters depth.
Stomiidae is a family of deep-sea ray-finned fish, including the barbeled dragonfishes. They are quite small, usually around 15 cm, up to 26 cm. These fish are apex predators and have enormous jaws filled with fang-like teeth. They are also able to hinge the neurocranium and upper-jaw system, which leads to the opening of the jaw to more than 100 degrees. This ability allows them to consume extremely large prey, often 50% greater than their standard length.
Malacosteus niger, commonly known as the black dragon fish, is a species of deep-sea fish. Some additional common names for this species include: northern stoplight loosejaw, lightless loosejaw, black loosejaw, and black hinged-head. It belongs to the family Stomiidae, or dragonfishes. It is among the top predators of the open mesopelagic zone. M. niger is a circumglobal species, which means that it inhabits waters ranging from the tropics to the subarctics. Not many studies have been conducted on its feeding habits, but recent research suggests that M. niger primarily feed on calanoid copepods which is a form of zooplankton. Indeed, it appears that M. niger primarily prey on zooplankton despite its apparent morphological adaptations for the consumption of relatively large prey. Another unique adaptation for this species is its ability to produce both red and blue bioluminescence. Most mesopelagic species aren't capable of producing red bioluminescence. This is advantageous because most other species cannot perceive red light, therefore allowing M. niger to camouflage part of itself to its prey and predators.
Sloane's viperfish, Chauliodus sloani, is a predatory mesopelagic dragonfish found in waters across the world. The species was first described by German scientists Marcus Elieser Bloch and Johann Gottlob Schneider in their 1801 book Systema ichthyologiae: iconibus CX illustratum, volume 1. Female C. sloani reach maturity between 133 and 191 mm, while males likely reach maturity at slightly smaller body lengths. It has two rows of photophores along its ventral side. It is believed that C. sloani can adjust the intensity of bioluminescence of the ventral photophores to camouflage itself from predators that might see its shadow from below.
The blackbelly lanternshark or lucifer shark is a shark of the family Etmopteridae found around the world in tropical and temperate seas at depths between 150 and 1,250 meters – the mesopelagic zone. Compared to other mesopelagic fish predators and invertebrates, the blackbelly lanternshark is thought to reside in shallower, more southern waters. E. lucifer can reach up to 47 centimeters in length and consumes mesopelagic cephalopods, fish, and crustaceans. Blackbelly lanternsharks are bioluminescent, using hormone controlled mechanisms to emit light through ventral photogenic organs called photophores and are presumed to be ovoviviparous. The blackbelly lanternshark has been classified as "Not Threatened" within the New Zealand Threat Classification System.
The viper dogfish or viper shark is a rare species of dogfish shark in the family Etmopteridae, and the only extant member of its genus. It has been found in the Pacific Ocean off southern Japan, the Bonin Islands, Pacific Ocean off northern Taitung County and the Northwestern Hawaiian Islands. This species inhabits upper continental slopes and seamounts. It may migrate vertically, shifting between bottom waters 270–360 m (890–1,180 ft) deep during the day and upper waters less than 150 m (490 ft) deep at night. A slender, black shark reaching 54 cm (21 in) in length, the viper dogfish can be recognized by its narrow, triangular jaws and well-spaced, fang-like teeth. It also has two spined dorsal fins, dermal denticles with faceted crowns, and numerous light-emitting photophores concentrated on its ventral surface.
The velvet belly lanternshark is a species of dogfish shark in the family Etmopteridae. One of the most common deepwater sharks in the northeastern Atlantic Ocean, the velvet belly is found from Iceland and Norway to Gabon and South Africa at a depth of 20–2,490 m (66–8,169 ft). A small shark generally no more than 45 cm (18 in) long, the velvet belly is so named because its black underside is abruptly distinct from the brown coloration on the rest of its body. The body of this species is fairly stout, with a moderately long snout and tail, and very small gill slits. Like other lanternsharks, the velvet belly is bioluminescent, with light-emitting photophores forming a species-specific pattern over its flanks and abdomen. The ventral photophores are thought to function in counter-illumination, which camouflages the shark against predators and prey. The bioluminescent flank markings may play a role in intraspecific communication.
The splendid lanternshark is a shark of the family Etmopteridae found in the western Pacific at depths between 120 and 210 m. Through the classification of Etmopterus species into several clades based on the positioning of their bioluminescent photophores, the splendid lanternshark can be considered a member of the Etmopterus pusillus clade.
Malacosteus australis, the southern stoplight loosejaw, is a species of barbeled dragonfish. This species is mainly distinguished from Malacosteus niger by a smaller postorbital photophore in both sexes and lower numbers of lateral photophores. It also differs in having somewhat smaller jaws, a fleshy orbit, and several subtle morphological traits. The maximum known length is 253.2 mm. Its specific epithet comes from the Latin austral, meaning "southern". It is known for its red bioluminescence which helps M. australis visualize in the aphotic deep sea.
Aristostomias is a genus of barbeled dragonfishes native to the ocean depths in the Pacific, Atlantic and Indian oceans.
Pachystomias microdon, the smalltooth dragonfish, is a species of barbeled dragonfish found in the oceans at depths of from 660 to 4,000 metres. This species grows to a length of 22.1 centimetres (8.7 in) SL. This species is the only known species in its genus.
Stomiati is a group of teleost fish belonging to the cohort (group) Euteleostei, which is a group of bony fishes within the infra-class Teleostei that evolved ~240 million years ago. Teleostei is a group of ray-finned fishes with the exception of primitive bichirs, sturgeons, paddlefishes, freshwater garfishes, and bowfins. The cohort of Euteleostei is divided into two smaller groups: the Protacanthopterygii and the Neoteleostei. Stomiati happen to be descendants of the Protacanthopterygii, and contains the order of Osmeriformes and Stomiiformes.
Peter John Herring is an English marine biologist known for his work on the coloration, camouflage and bioluminescence of animals in the deep sea, and for the textbook The Biology of the Deep Ocean.