This article possibly contains original research .(May 2019) |
Barreleyes | |
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Opisthoproctus soleatus . This image is drawn from a specimen taken to the surface; in a live specimen, the membrane over the top of the head forms a transparent dome. | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Argentiniformes |
Family: | Opisthoproctidae Schmidt, 1918 |
Genera | |
Barreleyes, also known as spook fish (a name also applied to several species of chimaera), are small deep-sea argentiniform fish comprising the family Opisthoproctidae found in tropical-to-temperate waters of the Atlantic, Pacific, and Indian Oceans. [2] [3] [4] [5]
These fish are named because of their barrel-shaped, tubular eyes, which are generally directed upwards to detect the silhouettes of available prey; however, the fish are capable of directing their eyes forward, as well. The family name Opisthoproctidae is derived from the Greek words opisthe 'behind' and proktos 'anus'.
The morphology of the Opisthoproctidae varies between three main forms: the stout, deep-bodied barreleyes of the genera Opisthoproctus and Macropinna , the extremely slender and elongated spookfishes of the genera Dolichopteryx and Bathylychnops , and the intermediate fusiform spookfishes of the genera Rhynchohyalus and Winteria .
All species have large, telescoping eyes, which dominate and protrude from the head, but are enclosed within a large transparent dome of soft tissue. [6] These eyes generally gaze upwards, but can also be directed forwards. [7] The opisthoproctid eye has a large lens and a retina with an exceptionally high complement of rod cells and a high density of rhodopsin (the "visual purple" pigment); no cone cells are present. To better serve their vision, barreleyes have large, dome-shaped, transparent heads; this presumably allows the eyes to collect even more incident light and likely protects the sensitive eyes from the nematocysts (stinging cells) of the siphonophores, from which the barreleye is believed to steal food. [8] It may also serve as an accessory lens (modulated by intrinsic or peripheral muscles), or refract light with an index very close to seawater. Dolichopteryx longipes is the only vertebrate known to use a mirror (as well as a lens) in its eyes for focusing images. [9]
The toothless mouth is small and terminal, ending in a pointed snout. As in related families (e.g. Argentinidae), an epibranchial or crumenal organ is present behind the fourth gill arch. This organ—analogous to the gizzard—consists of a small diverticulum wherein the gill rakers insert and interdigitate for the purpose of grinding up ingested material. The living body of most species is a dark brown, covered in large, silvery imbricate scales, but these are absent in Dolichopteryx, leaving the body itself a transparent white. In all species, a variable number of dark melanophores colour the muzzle, ventral surface, and midline.
Also present in Dolichopteryx, Opisthoproctus, and Winteria species are a number of luminous organs; Dolichopteryx has several along the length of its belly, and Opisthoproctus has a single organ in the form of a rectal pouch. These organs glow with a weak light due to the presence of symbiotic bioluminescent bacteria, specifically, Photobacterium phosphoreum (family Vibrionaceae). The ventral surfaces of Opisthoproctus species are characterised by a flattened and projecting 'sole'; in the mirrorbelly (Opisthoproctus grimaldii) and Opisthoproctus soleatus , this sole may act as a reflector, by directing the emitted light downwards. The strains of P. phosphoreum present in the two Opisthoproctus species have been isolated and cultured in the lab. Through restriction fragment length polymorphism analysis, the two strains have been shown to differ only slightly. [10] [11]
In all species, the fins are spineless and fairly small; in Dolichopteryx however, the pectoral fins are greatly elongated and wing-like, extending about half the body's length, and are apparently used for stationkeeping in the water column. The pectoral fins are inserted low on the body, and in some species, the pelvic fins are inserted ventrolaterally rather than strictly ventrally. Several species also possess either a ventral or dorsal adipose fin, and the caudal fin is forked to emarginated. The anal fin is either present or greatly reduced, and may not be externally visible; it is strongly retrorse in Opisthoproctus. A single dorsal fin originates slightly before or directly over the anal fin. A perceptible hump in the back begins just behind the head. The gas bladder is absent in most species, and the lateral line is uninterrupted. The branchiostegal rays (bony rays supporting the gill membranes behind the lower jaw) number two to four. The javelin spookfish (Bathylychnops exilis) is by far the largest species at 50 centimetres (20 in) standard length; most other species are under 20 centimetres (7.9 in).
Barreleyes inhabit moderate depths, from the mesopelagic to bathypelagic zone, circa 400–2,500 m deep. They are presumably solitary and do not undergo diel vertical migrations; instead, barreleyes remain just below the limit of light penetration and use their sensitive, upward-pointing tubular eyes—adapted for enhanced binocular vision at the expense of lateral vision—to survey the waters above. The high number of rods in their eyes' retinae allows barreleyes to resolve the silhouettes of objects overhead in the faintest of ambient light (and to accurately distinguish bioluminescent light from ambient light), and their binocular vision allows the fish to accurately track and home in on small zooplankton such as hydroids, copepods, and other pelagic crustaceans. The distribution of some species coincides with the isohaline and isotherm layers of the ocean; for example, in Opisthoproctus soleatus, upper distribution limits coincide with the 400-m isotherm for 8 °C (46 °F).
What little is known of barreleye reproduction indicates they are pelagic spawners; that is, eggs and sperm are released en masse directly into the water. The fertilized eggs are buoyant and planktonic; the larvae and juveniles drift with the currents—likely at much shallower depths than the adults—and upon metamorphosis into adult form, they descend to deeper waters. Dolichopteryx species are noted for their paedomorphic features, the result of neoteny (the retention of larval characteristics).
The bioluminescent organs of Dolichopteryx and Opisthoproctus, together with the reflective soles of the latter, may serve as camouflage in the form of counterillumination . This predator avoidance strategy involves the use of ventral light to break up the fishes' silhouettes, so that (when viewed from below) they blend in with the ambient light from above. Counterillumination is also seen in several other unrelated deep-sea families, which include the marine hatchetfish (Sternoptychidae). Also found in marine hatchetfish and other unrelated families are tubular eyes, such as telescopefish and tube-eye.
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.
Barracudinas are any member of the marine mesopelagic fish family Paralepididae: 50 or so extant species are found almost worldwide in deep waters. Several genera are known only from fossils dating back to the Ypresian epoch.
Marine hatchetfishes or deep-sea hatchetfishes are small deep-sea mesopelagic ray-finned fish of the stomiiform subfamily Sternoptychinae. They should not be confused with the freshwater hatchetfishes, which are not particularly closely related Teleostei in the characiform family Gasteropelecidae.
The marine hatchetfishes or deep-sea hatchetfishes as well as the related bottlelights, pearlsides and constellationfishes are small deep-sea ray-finned fish of the stomiiform family Sternoptychidae. They are not closely related to and should not be confused with the freshwater hatchetfishes, which are teleosts in the characiform family Gasteropelecidae. The Sternoptychidae have 10 genera and about 70 species altogether.
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.
Sabertooth or sabretooth fish are small, fierce-looking deep-sea aulopiform fish comprising the family Evermannellidae. The family is small, with just eight species in three genera represented; they are distributed throughout tropical to subtropical waters of the Atlantic, Indian, and Pacific Oceans.
Telescopefish are small, deep-sea aulopiform fish comprising the small family Giganturidae. The two known species are within the genus Gigantura. Though rarely captured, they are found in cold, deep tropical to subtropical waters worldwide.
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.
Photobacterium phosphoreum is a Gram-negative, bioluminescent bacterium living in symbiosis with deep-sea marine organisms, such as anglerfish. It can emit bluish-green light (490 nm) due to a chemical reaction between FMN, luciferin and molecular oxygen catalysed by an enzyme called luciferase.
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.
Monacoa grimaldii, also known as the mirrorbelly, barreleye, Grimaldi's barreleye, and flatiron fish, is a species of fish in the family Opisthoproctidae. Different sources express different views on its distribution. According to Poulsen and colleagues, it is known with certainty only from the Atlantic Ocean, with records from the Pacific Ocean representing other Monacoa species. However, FishBase and Catalog of Fishes include the Pacific, and in the case of the latter, the Indian Ocean in its range. It is typically mesopelagic, but it has been recorded from depths of 0 to 4,750 metres. It has a body length of 8 cm (3.1 in) SL.
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 brownsnout spookfish or brown-snout spookfish is a species of barreleye in the family Opisthoproctidae. It and the glasshead barreleye fish are the only vertebrates known to employ a mirror, in addition to a lens, to focus an image in its eyes. This species probably has a worldwide tropical and temperate distribution; in the Atlantic Ocean it is known from Bermuda, the Bahamas, the Greater Antilles, and the Gulf of Mexico, and in the Pacific Ocean it is known from the California Current region and the South China Sea. It is found in the mesopelagic and bathypelagic zones at a depth of 500–2,400 meters (1,600–7,900 ft), but usually occurs below 1,000 meters. In the Gulf of Mexico it is found shallower, at 310–460 meters (1,020–1,510 ft).
The Pacific barreleye fish (Macropinna) is a genus of ray-finned fish belonging to Opisthoproctidae, the barreleye family. It contains one species, M. microstoma. It is recognized for a highly unusual transparent, fluid-filled shield on its head, through which the lenses of its eyes can be seen. It was originally believed that the tubular eyes of this fish were fixed in place and, therefore, only provided a tunnel vision view of what was seen above its head. However, in 2008, scientists discovered that its eyes were able to rotate both up and forward in its transparent shield. M. microstoma has a tiny mouth and most of its body is covered with large scales. The fish normally hangs nearly motionless in the water, at a depth of about 600 metres (2,000 ft) to 800 metres (2,600 ft), using its large fins for stability and with its eyes directed upward. In the low light conditions it is assumed the fish detects prey by its silhouette. MBARI researchers Bruce Robison and Kim Reisenbichler observed that when prey such as small fish and jellyfish are spotted, the eyes rotate like binoculars, facing forward as it turns its body from a horizontal to a vertical position to feed. Robison speculates that M. microstoma steals food from siphonophores.
Underwater camouflage is the set of methods of achieving crypsis—avoidance of observation—that allows otherwise visible aquatic organisms to remain unnoticed by other organisms such as predators or prey.
Vision is an important sensory system for most species of fish. Fish eyes are similar to the eyes of terrestrial vertebrates like birds and mammals, but have a more spherical lens. Birds and mammals normally adjust focus by changing the shape of their lens, but fish normally adjust focus by moving the lens closer to or further from the retina. Fish retinas generally have both rod cells and cone cells, and most species have colour vision. Some fish can see ultraviolet and some are sensitive to polarised light.
Opisthoproctus soleatus is a species of fish in the family Opisthoproctidae. It was first described in 1888 by Léon Vaillant. The species lives in most tropical seas, but is more common in the eastern Atlantic, from western Ireland to Mauritania and from Sierra Leone to Angola, and also in the South China Sea. O. soleatus can grow to a standard length of 10.5 centimetres (4.1 in) and usually live from about 500 to 700 metres deep.
Argyropelecus affinis is a species of ray-finned fish in the family Sternoptychidae, found in the tropical and subtropical Atlantic, Indian and Pacific Oceans. Common names for this fish include Pacific hatchetfish, deepsea hatchetfish and slender hatchetfish. It inhabits the mesopelagic zone and is either non-migratory or performs short daily vertical migrations.
Neoscopelus macrolepidotus, also known as a large-scaled lantern fish, is a species of small mesopelagic or bathypelagic fish of the family Neoscopelidae, which contains six species total along three genera. The family Neoscopelidae is one of the two families of the order Myctophiformes. Neoscopelidae can be classified by the presence of an adipose fin. The presence of photophores, or light-producing organs, further classify the species into the genus Neoscopelus. N. macrolepidotus tends to be mesopelagic until the individuals become large adults, which is when they settle down to the bathypelagic zone.
Argyropelecus olfersii is a common species of marine hatchetfish, found in mesopelagic waters.