Malacosteus niger | |
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Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Actinopterygii |
Order: | Stomiiformes |
Family: | Stomiidae |
Genus: | Malacosteus |
Species: | M. niger |
Binomial name | |
Malacosteus niger Ayres, 1848 | |
Synonyms | |
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. [1] It belongs to the family Stomiidae, or dragonfishes. It is among the top predators of the open mesopelagic zone. [2] M. niger is a circumglobal species, which means that it inhabits waters ranging from the tropics to the subarctics. [3] 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. [2] 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.
Malacosteus niger has yellow lenses that are believed to improve the functionality of the perception of their red bioluminescence. M. niger has adapted a retinal structure of "ten layer elements," similar to those found in surface-level species and other shallow-water living species — which also perceive red light. [4] Its retina is made up entirely of rods and no cones, with rhodopsin/porphyropsin pairs and a single opsin bound to some of its photoreceptors, which provide visual sensitivity up to 517-541 nm (this falls within the wavelength of red light). [5] Most deep-sea fish have a single visual pigment maximally sensitive at short wavelengths, approximately matching the spectrum of both downwelling sunlight and bioluminescence. [6] For comparison, other red light producing stomiids, such as Aristostomias and Pachystostomias, have a third pigment which allows them to perceive light up to 588 nm and 595 nm respectively. The yellow lens reduce the amount of blue light that reaches the retina and increases sensitivity to longer wavelengths, which benefits M. niger and its red bioluminescence.
Yellow lens have also been identified in Echiostoma , which also produces red bioluminescence.
Malacosteus niger has one of the largest relative gapes of any fish with the lower jaw being approximately one-quarter of the fishes length (Figure A). [2] It has enlarged fangs the curve back into its mouth to prevent its prey from escaping its grip (Figure B). M. nigers is unique in that it does not contain gill rakers or gill teeth which is typically found in carnivorious fish species (Figure C). The anterior vertebrae appear to be unossified which enables the fish to “throw back its head” to take on relatively large prey. [7] Lastly, M. nigers lacks an ethinoid membrane (no “floor” in its mouth) which allows for it to consume bigger prey species (Figure D). [2] (Refer to image on right hand side of webpage). The lack of a floor of the oral cavity allows for decreased resistive forces which allows M. niger to close its mouth rapidly and easily trap its prey. This adaptation also minimizes the amount of energy required for M. niger to close its mouth, thus permitting it to quickly latch onto fast-swimming prey. [8]
The postorbital photophore in this species is larger than in M. australis . It also differs in lateral photophore count, as well as in morphological characters. The maximum known length is 25.6 cm (10.1 in). Its specific epithet niger is Latin for "black". [9]
Malacosteus niger is a circumglobal species and has a large geographic range. It can typically be found from the Arctic latitudes of 66° North and from 30° South in the Southern Hemisphere. [2] While M. nigers is found throughout the world, it appears to be widely distributed and found often in the Eastern Central Atlantic Ocean. [10] M. niger does not leave the mesopalgic and is known to have a vertical range from 500 meters to 1000 meters. [1] Interestingly, it is believed to be the only member of the family that does not go through diel/vertical migration, which means that it does not migrate up to the surface like other fish species. [9]
While the morphology of M. niger with huge fangs and an enormous gape is typical for its family and suggests adaptations to piscivory, its diet in fact contains a substantial proportion of zooplankton. [11] Some of its documented prey include calanoid copepods, micronekton, decapod shrimps, and other decapods. [2] M. niger digests its prey within a diel cycle, meaning the copepods it consumes at nighttime are digested by the afternoon the following day, which requires it to be constantly feeding on these small prey to sustain its energy. It has been recorded that copepods make up around 69%-83% of M. niger's diet. [11] This suggests that availability of large prey at these depths is often limited. It is suggested that its dominant feeding mode is searching for zooplanktonic prey (copepods in particular) using bioluminescence to illuminate a small search area, since M. niger experiences infrequent encounters with larger prey items. Current research suggests that M. niger has adopted this unique feeding habit in association with the abundance of prey. Recent studies in the eartern Gulf of Mexico indicate that large calanoid copepods are three orders of magnitude more abundant than either fish or shrimp. [11] More research is needed within different regions to confirm this hypothesis. It is suggested that its dominant feeding mode is searching for zooplanktonic prey (copepods in particular) using bioluminescence to illuminate a small search area, with infrequent encounters with larger prey items. The likely origin of the pigment necessary for detecting its long wavelength bioluminescence, a chlorophyll derivative, is the copepods themselves. [11] [12]
Malacosteus niger's unique adaptation of producing red bioluminescence is only found in two other deep-sea dwelling creatures, Aristostomias and Pachystomias. [13] This rare form of bioluminescence can reach up to 700 nm in the deep-sea and cannot be perceived by green and blue bioluminescent organisms, thus granting M. niger a considerable advantage while hunting for food. [5] Other deep-sea fish capable of detecting far-red bioluminescence, including Aristostomias and Pachystomias are able to do so using visual pigments. M. niger lacks these same long-wave pigments, and instead increases its sensitivity to red light using a chlorophyll-derived photosensitizer. [14]
Malacosteus Niger has a tear-drop shaped, dark brown, suborbital photophore which is used to emit red light at an emission maxima of 710 nanometers. Removal of the top brown photophore layer causes a shift of the emission spectra to shorter wavelengths of around 650 nanometers. The photophores contain red-fluorescent material which is made to fluoresce via energy transfer from chemical reactions. Control of the photophore is maintained via innervation through branches of the fifth cranial nerve and this photophore. It is reported to be controlled independently of the postorbital blue photophore and has been noted to fluoresce for longer durations. The photophore is composed of a large pigmented sac containing a mass of scarlet gland cells. A thick, reflective layer lines the pigment sac, with occasional strands of reflective tissue running through the glandular core of the photophore. The outer layer is composed of large epithelial cells which merge into an inner, darker stained layer. The presumed function of this layer is to provide the brown layer through which fluorescence is filtered. The cells of the glandular core are characterized by a dense rough endoplasmic reticulum. [15]
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.
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.
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. This genus once contained three nominal species: M. niger, 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.
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 firefly squid, also commonly known as the sparkling enope squid or hotaru-ika in Japan, is a species of squid in the family Enoploteuthidae. W. scintillans is the sole species in the monotypic genus Watasenia.
Biological pigments, also known simply as pigments or biochromes, are substances produced by living organisms that have a color resulting from selective color absorption. Biological pigments include plant pigments and flower pigments. Many biological structures, such as skin, eyes, feathers, fur and hair contain pigments such as melanin in specialized cells called chromatophores. In some species, pigments accrue over very long periods during an individual's lifespan.
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. 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 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.
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.
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.