Stomiiformes is an
order
Order, ORDER or Orders may refer to:
* Categorization, the process in which ideas and objects are recognized, differentiated, and understood
* Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...
of
deep-sea ray-finned fish
Actinopterygii (; ), members of which are known as ray-finned fishes, is a class of bony fish. They comprise over 50% of living vertebrate species.
The ray-finned fishes are so called because their fins are webs of skin supported by bony or h ...
es of very diverse
morphology. It includes, for example,
dragonfishes, lightfishes (
Gonostomatidae and
Phosichthyidae),
loosejaw
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 ...
s,
marine hatchetfishes and
viperfishes. The order contains 4
families (5 according to some authors) with more than 50
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 ...
and at least 410
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 ...
. 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.
The
scientific name means "''Stomias''-shaped", from ''
Stomias'' (the
type genus
In biological taxonomy, the type genus is the genus which defines a biological family and the root of the family name.
Zoological nomenclature
According to the International Code of Zoological Nomenclature, "The name-bearing type of a nomina ...
) + the standard
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% ...
order
suffix "-formes". It ultimately derives from
Ancient Greek
Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic p ...
''stóma'' (στόμᾶ, "mouth") +
Latin
Latin (, or , ) is a classical language belonging to the Italic branch of the Indo-European languages. Latin was originally a dialect spoken in the lower Tiber area (then known as Latium) around present-day Rome, but through the power ...
''forma'' ("external form"), the former in reference to the huge mouth opening of these fishes.
Description and ecology
Members of this order are mostly
pelagic
The pelagic zone consists of the water column of the open ocean, and can be further divided into regions by depth (as illustrated on the right). The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or w ...
fishes living in deep oceanic waters. Their distribution around the world's oceans is very wide, ranging from
subtropical and
temperate
In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
waters up to
subarctic or even
Antarctic
The Antarctic ( or , American English also or ; commonly ) is a polar region around Earth's South Pole, opposite the Arctic region around the North Pole. The Antarctic comprises the continent of Antarctica, the Kerguelen Plateau and o ...
ones.
[FishBase (2005), Nelson (2006): p.207]

The smallest species of this order is the
bristlemouth ''
Cyclothone pygmaeae
''Cyclothone'' is a genus containing 13 extant species of Bioluminescence, bioluminescent fish, commonly known as 'bristlemouths' or 'bristlefishes' due to their shared characteristic of sharp, bristle-like teeth. These fishes typically grow to ...
''. Native to the
Mediterranean Sea
The Mediterranean Sea is a sea connected to the Atlantic Ocean, surrounded by the Mediterranean Basin and almost completely enclosed by land: on the north by Western and Southern Europe and Anatolia, on the south by North Africa, and on the ...
, it reaches just 1.5 cm (0.6 in) as an adult. The largest species is the
barbeled dragonfish ''
Opostomias micripnis'', widely found in the
Atlantic
The Atlantic Ocean is the second-largest of the world's five oceans, with an area of about . It covers approximately 20% of Earth's surface and about 29% of its water surface area. It is known to separate the "Old World" of Africa, Europe an ...
,
Indian and
Pacific Ocean
The Pacific Ocean is the largest and deepest of Earth's five oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean (or, depending on definition, to Antarctica) in the south, and is bounded by the contine ...
s and measuring about in adult length.
These fish have a highly unusual and often almost nightmarish appearance. They all have teeth on the
premaxilla
The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammal has ...
and
maxilla
The maxilla (plural: ''maxillae'' ) in vertebrates is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The ...
. Their maxillary
ligaments, as well as some muscles and certain bones in the
branchial cavity Branchial may refer to:
* Branchial apparatus, an embryological structure.
* Branchial arch a series of bony "loops" present in fish, which support the gills.
* Branchial artery, also known as aortic arches.
* Branchial cleft
* Branchial cleft ...
, are specialized in a distinctive way. Most have large mouths extending back past the eyes. Some also have a chin
barbel. The
dorsal
Dorsal (from Latin ''dorsum'' ‘back’) may refer to:
* Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism
* Dorsal, positioned on top of an aircraft's fuselage
* Dorsal co ...
and/or
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 ...
s are missing in some, but others have 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 s ...
. The
pelvic fin
Pelvic fins or ventral fins are paired fins located on the ventral surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods.
Structure and function Structure
In actinopterygians, the pelvic fin consists of two e ...
has 4-9
rays, and the stomiiformes possess 5-24
branchiostegal ray
This glossary of ichthyology is a list of definitions of terms and concepts used in ichthyology, the study of fishes.
A
B
C
...
s. Their
scale
Scale or scales may refer to:
Mathematics
* Scale (descriptive set theory), an object defined on a set of points
* Scale (ratio), the ratio of a linear dimension of a model to the corresponding dimension of the original
* Scale factor, a number ...
s are
cycloid
In geometry, a cycloid is the curve traced by a point on a circle as it rolls along a straight line without slipping. A cycloid is a specific form of trochoid and is an example of a roulette, a curve generated by a curve rolling on another ...
, delicate and easily sloughed off; some are scaleless. The coloration is typically dark brown or black; a few (mostly
Gonostomatoidei) are silver, and
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, h ...
s (light-producing organs) are common in this order.
The teeth of stomiiformes are often transparent and non-reflective so that prey will be unlikely to see them in the light generated by bioluminescence. Research has revealed that the transparency of the teeth of ''
Aristostomias scintillans'' is due to nanoscale structures composed
hydroxyapatite
Hydroxyapatite, also called hydroxylapatite (HA), is a naturally occurring mineral form of calcium apatite with the formula Ca5(PO4)3(OH), but it is usually written Ca10(PO4)6(OH)2 to denote that the crystal unit cell comprises two entities ...
and
collagen and a lack
dentin tubules, however a study from a decade prior had shown the teeth of ''
Chauliodus sloani'' (which are also transparent) have dentin tubules. The reason behind difference in presence of dentin tubules in two species of the same family (Stomiidae) has yet to be addressed.
Bioluminescence

As common for
deep-sea creatures, all members of Stomiiformes (except one) have
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, h ...
s, whose structure is characteristic of the order. The light emitted can be more or less strong and its color can be light yellow, white, violet or red. The light coming from these fish is generally invisible to their prey. The lighting mechanism can be very simple – consisting of small gleaming points on the fish body – or very elaborate, involving
lens
A lens is a transmissive optical device which focuses or disperses a light beam by means of refraction. A simple lens consists of a single piece of transparent material, while a compound lens consists of several simple lenses (''elements'' ...
es and
refractor
A refracting telescope (also called a refractor) is a type of optical telescope that uses a lens as its objective to form an image (also referred to a dioptric telescope). The refracting telescope design was originally used in spyglasses and ...
s.
The most common arrangement is one or two rows of photophores on the ventral aspect of the body. The rows run from the head down to the tip of the tail. Photophores are also present in chin barbels of the family
Stomiidae. The light produced in these glandular organs is the product of an enzymatic reaction, a catylization of
coelenterazine by calcium ions.
Daily migration
During the day, Stomiiformes stay in deep waters. When the sun sets, most of them follow the dimming sunlight up to near-surface waters, which are richer in animal life such as small fishes and
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 cr ...
ic
invertebrate
Invertebrates are a paraphyletic group of animals that neither possess nor develop a vertebral column (commonly known as a ''backbone'' or ''spine''), derived from the notochord. This is a grouping including all animals apart from the chordate ...
s. During the night, these Stomiiformes hunt and feed on such organisms, swimming back to deeper waters when the sun rises. They apparently are able to measure the intensity of the sunlight that reaches them. They will thus move to stay always in the zone where
light intensity is very low, though it is not entirely dark.
This daily migration is well observed in quite a few species of stomiiforms. However, it is also performed by other fishes, while some larger Stomiiformes – among them the largest
predator
Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill ...
s of the deep sea – stay in their
habitat
In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
all the time and feed on smaller migrating fish that return from the surface.
Reproduction
Stomiiforms
spawn generally in deep seas, but the eggs are light and float towards the ocean surface. They hatch in surface waters. When the
larva
A larva (; plural larvae ) is a distinct juvenile form many animals undergo before metamorphosis into adults. Animals with indirect development such as insects, amphibians, or cnidarians typically have a larval phase of their life cycle.
Th ...
e have completed their
metamorphosis and look like adults, they descend to join the main population.
Like many
benthic
The benthic zone is the ecological region at the lowest level of a body of water such as an ocean, lake, or stream, including the sediment surface and some sub-surface layers. The name comes from ancient Greek, βένθος (bénthos), meaning "t ...
fish species, certain members of the order – especially in the genera ''
Cyclothone
''Cyclothone'' is a genus containing 13 extant species of bioluminescent fish, commonly known as 'bristlemouths' or 'bristlefishes' due to their shared characteristic of sharp, bristle-like teeth. These fishes typically grow to around 1-3 inches ...
'' and ''
Gonostoma'' – change their sex during their life. When they become
sexually mature
Sexual maturity is the capability of an organism to reproduce. In humans it might be considered synonymous with adulthood, but here puberty is the name for the process of biological sexual maturation, while adulthood is based on cultural defini ...
, they are males; later on they transform into females.
Systematics
The Stomiiformes are often placed in the
teleost
Teleostei (; Greek ''teleios'' "complete" + ''osteon'' "bone"), members of which are known as teleosts ), is, by far, the largest infraclass in the class Actinopterygii, the ray-finned fishes, containing 96% of all extant species of fish. Tel ...
superorder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
Stenopterygii, usually together with the
Ateleopodiformes (jellynoses), but sometimes on their own. Whether it is indeed justified to accept such a small group is doubtful; it may well be that the closest living relatives of the "Stenopterygii" are found among the superorder
Protacanthopterygii, and that the former would need to be merged in the latter. In some classifications, the "Stenopterygii" are kept separate but included with the Protacanthopterygii and the
monotypic
In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...
superorder
Cyclosquamata in an unranked
clade called Euteleostei. That would probably require splitting two additional monotypic superorders out of the Protacanthopterygii, and thus result in a profusion of very small
taxa
In biology, a taxon ( back-formation from '' taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular n ...
.

The Stomiiformes have also been considered close relatives of the
Aulopiformes
Aulopiformes is a diverse order of marine ray-finned fish consisting of some 15 extant and several prehistoric families with about 45 genera and over 230 species. The common names grinners, lizardfishes and allies, or aulopiforms are sometimes ...
. The latter are otherwise placed in a monotypic superorder "Cyclosquamata" but also appear to be quite close to the Protacanthopterygii indeed. The relationships of these – and the
Lampriformes or
Myctophiformes
The Myctophiformes are an order of ray-finned fishes consisting of two families of deep-sea marine fish, most notably the highly abundant lanternfishes (Myctophidae). The blackchins (Neoscopelidae) contain six species in three genera, while the ...
, which are also usually treated as monotypic superorders – to the taxa mentioned before is still not well resolved at all, and regardless whether one calls them Protacanthopterygii ''
sensu lato
''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
'' or Euteleostei, the
phylogeny
A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spe ...
of this group of moderately-advanced Teleostei is in need of further study.
[FishBase (2005), Diogo (2008)]
The ancestral Stomiiformes probably had thin brownish bodies, rows of egg-shaped
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, h ...
s adorning the lower body parts, and mouths with numerous teeth. From these, two lineages
evolved
Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation te ...
, probably some time during the
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the younger of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''creta'', ...
: Among the modern Stomiiformes, the
Gonostomatidae and
Phosichthyidae are
phenetically very similar, but this is due to their being very
plesiomorphic and retaining many traits of the original stomiiforms. Each of the two has characteristic
synapomorph
In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to have ...
ies with one of the more advanced stomiiform families – the
Sternoptychidae and the
Stomiidae, respectively. These two, in turn, are highly
autapomorphic, and at a casual glance do not look as if they were as closely related to the other stomiiforms as they actually are.
Thus, the classification of the
suborder
Order ( la, ordo) is one of the eight major hierarchical taxonomic ranks in Linnaean taxonomy. It is classified between family and class. In biological classification, the order is a taxonomic rank used in the classification of organisms and ...
s and
families of the Stomiiformes is:
* Suborder
Gonostomatoidei
** Family
Gonostomatidae – bristlemouths, anglemouths, "lightfishes" (including Diplophidae)
** Family
Sternoptychidae – marine hatchetfishes, bottlelights, constellationfishes, pearlsides
* Suborder
Stomioidei
** Family
Phosichthyidae – lightfishes
** Family
Stomiidae – barbeled dragonfishes, loosejaws, stareaters
Timeline of genera
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** Early Christianity
** Early modern Europe
Places in the United States
* Early, Iowa
* Early, Texas
* Early ...
from: -99.6 till: -65.5 color:latecretaceous text: Late
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from: -5.332 till: -2.588 color:pliocene text:Pl.
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* Liberal Party (Brazil, 2006), a Brazilian political party
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* Liberal Party (Brazil, 2006), a Brazilian political party
* Liberal Party (Moldova), a Moldovan political p ...
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The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
from: -65.5 till: -23.03 color:paleogene text:Paleogene
The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago (Mya) to the beginning of ...
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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. ...
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color:latecretaceous bar:NAM1 from:-99.6 till:-97.6 text: Paravinciguerria
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*Idrissa Camará (born 1992), Bissau-Guinean footballer
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color:eocene bar:NAM3 from:-55.8 till:0 text: Polymetme
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''Valenciennellus'' is a genus of marine hatchetfishes. They are commonly known as constellationfishes.
Species
There are currently two recognized species in this genus:
* ''Valenciennellus carlsbergi'' Bruun, 1931
* ''Valenciennellus tripunct ...
color:eocene bar:NAM5 from:-55.8 till:0 text:Argyropelecus
''Argyropelecus'' is an oceanic ray-finned fish genus in the deep sea hatchetfish family Sternoptychidae. A collective name is " silver hatchetfishes", but this can also refer to a species of the freshwater hatchetfishes which are not particula ...
color:eocene bar:NAM6 from:-48.6 till:-37.2 text: Polyipnoides
color:eocene bar:NAM7 from:-48.6 till:0 text: Astronesthes
color:eocene bar:NAM8 from:-48.6 till:0 text: Vinciguerria
color:eocene bar:NAM9 from:-37.2 till:0 text:Danaphos
''Danaphos'' is an oceanic ray-finned fish genus which belongs in the family Sternoptychidae. A common name is bottlelights.
Species
There are currently two recognized species in this genus:
* ''Danaphos asteroscopus'' Bruun, 1931
* ''Dan ...
color:oligocene bar:NAM10 from:-33.9 till:-28.4 text: Praewoodsia
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''Polyipnus'' is a genus of oceanic ray-finned fish in the family Sternoptychidae. This is the largest genus of the marine hatchetfishes subfamily Sternoptychinae and indeed of the entire Sternoptychidae. It is not quite as apomorphic as their re ...
color:oligocene bar:NAM13 from:-33.9 till:0 text:Sternoptyx
''Sternoptyx'' is an oceanic ray-finned fish genus which belongs in the family Sternoptychidae. This is the type genus of the Sternoptychidae, as well as the marine hatchetfish subfamily Sternoptychinae.
''Sternoptyx'' have silvery, high ...
color:oligocene bar:NAM14 from:-28.4 till:0 text: Argyripnus
color:miocene bar:NAM15 from:-23.03 till:0 text: Polymetmeglareosus
color:miocene bar:NAM16 from:-23.03 till:0 text: Maurolicus
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color:miocene bar:NAM19 from:-11.61 till:0 text: Chauliodus
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color:miocene bar:NAM21 from:-11.61 till:0 text: Gonostoma
color:miocene bar:NAM22 from:-11.61 till:0 text: Ichthyococcus
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** Early Christianity
** Early modern Europe
Places in the United States
* Early, Iowa
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* Liberal Party (Brazil, 2006), a Brazilian political party
* Liberal Party (Moldova), a Moldovan political p ...
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* Liberal Party (Brazil, 2006), a Brazilian political party
* Liberal Party (Moldova), a Moldovan political p ...
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bar:era
from: -145.5 till: -65.5 color:cretaceous text:Cretaceous
The Cretaceous ( ) is a geological period that lasted from about 145 to 66 million years ago (Mya). It is the third and final period of the Mesozoic Era, as well as the longest. At around 79 million years, it is the longest geological period of ...
from: -65.5 till: -23.03 color:paleogene text:Paleogene
The Paleogene ( ; also spelled Palaeogene or Palæogene; informally Lower Tertiary or Early Tertiary) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period million years ago (Mya) to the beginning of ...
from: -23.03 till: -2.588 color:neogene text: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. ...
from: -2.588 till: 0 color:quaternary text: Q.
Footnotes
References
* (2008): On the cephalic and pectoral girdle muscles of the deep sea fish ''Alepocephalus rostratus'', with comments on the functional morphology and phylogenetic relationships of the Alepocephaloidei (Teleostei). ''
Anim. Biol.'' 58(1): 23-29.
* (2005)
Order Stomiiformes Version of 2005-FEB-15. Retrieved 2009-SEP-28.
*
* (1968-1982): ''
Oxford Latin Dictionary
The ''Oxford Latin Dictionary'' (or ''OLD'') is the standard English lexicon of Classical Latin, compiled from sources written before AD 200. Begun in 1933, it was published in fascicles between 1968 and 1982; a lightly revised second edition ...
'' (1st ed.). Oxford University Press, Oxford.
*
* (2006): ''
Fishes of the World
''Fishes of the World'' by the American Ichthyology, ichthyologist Joseph S. Nelson (1937–2011) is a standard reference for fish systematics. Now in its fifth edition (2016), the work is a comprehensive overview of the diversity and classificat ...
'' (4th ed.). John Wiley & Sons, Inc.
*
* (1910): ''English-Greek Dictionary - A Vocabulary of the Attic Language''. George Routledge & Sons Ltd., Broadway House, Ludgate Hill, E.C
Searchable JPEG fulltext*
{{Taxonbar, from=Q657570
Articles which contain graphical timelines
Ray-finned fish orders
Bioluminescent fish
Taxa named by Charles Tate Regan