The footballfish form a

family Family (from ) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). It forms the basis for social order. Ideally, families offer predictabili ...

, Himantolophidae, of globose,

deep-sea The deep sea is broadly defined as the ocean depth where light begins to fade, at an approximate depth of or the point of transition from continental shelves to continental slopes. Conditions within the deep sea are a combination of low tempe ...

anglerfish The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common name, common and scientific name comes from the characteristic mode of predation, in which a modified dorsal Fish fin#Ray-fins, fin ray acts as a Aggressiv ...

es found in tropical and subtropical waters of the

Atlantic The Atlantic Ocean is the second largest of the world's five oceanic divisions, with an area of about . It covers approximately 17% of Earth's surface and about 24% of its water surface area. During the Age of Discovery, it was known for se ...

Indian Indian or Indians may refer to: Associated with India * of or related to India ** Indian people ** Indian diaspora ** Languages of India ** Indian English, a dialect of the English language ** Indian cuisine Associated with indigenous peoples o ...

, and

Pacific Ocean The Pacific Ocean is the largest and deepest of Earth's five Borders of the oceans, oceanic divisions. It extends from the Arctic Ocean in the north to the Southern Ocean, or, depending on the definition, to Antarctica in the south, and is ...

. The family contains 23

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

, all of which are classified in a single

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

, ''Himantolophus''.

Etymology

The football fish family and genus names are derived from a combination of ''himantos'', which means a "leather strap", "thong" or "leash", with ''lophus'', meaning "crest" or "tuft". Reinhardt did not explain this name but it is thought to refer to the thick, leathery

illicium ''Illicium'' is a genus of flowering plants treated as part of the family Schisandraceae,

of the

type species In International_Code_of_Zoological_Nomenclature, zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the spe ...

, ''H. groenlandicus''.

Taxonomy

The footballfish genus, ''Himantolophus'' was first proposed as a

monospecific genus 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 "unispe ...

in 1837 by the Norwegian

zoologist Zoology ( , ) is the scientific study of animals. Its studies include the structure, embryology, classification, habits, and distribution of all animals, both living and extinct, and how they interact with their ecosystems. Zoology is one ...

Johan Reinhardt Johannes Christopher Hagemann Reinhardt (23 December 1778 – 31 October 1845), sometimes called J. C. H. Reinhardt, was a professor in zoology at the University of Copenhagen. Born in Rendal Municipality, Rendalen parish in Norway, his father, J ...

when he described ''Himantolophus groenlandicus''. Reinhardt gave the type locality of ''H. groenlandicus'' as being near

Godthaab Nuuk (; , formerly ) is the capital (political), capital and List of cities and towns in Greenland, most populous city of Greenland, an autonomous territory in the Kingdom of Denmark. Nuuk is the seat of Government of Greenland, government an ...

Greenland Greenland is an autonomous territory in the Danish Realm, Kingdom of Denmark. It is by far the largest geographically of three constituent parts of the kingdom; the other two are metropolitan Denmark and the Faroe Islands. Citizens of Greenlan ...

where it had been washed ashore. In 1861

Theodore Gill Theodore Nicholas Gill (March 21, 1837 – September 25, 1914) was an American ichthyologist, mammalogist, malacologist, and librarian. Career Born and educated in New York City under private tutors, Gill early showed interest in natural hist ...

placed ''Himantolophus'' in the new

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 "unisp ...

family Himantolophidae. The 5th edition of

Fishes of the World ''Fishes of the World'' is a standard reference for the systematics of fishes. It was first written in 1976 by the American ichthyologist Joseph S. Nelson (1937–2011). Now in its fifth edition (2016), the work is a comprehensive overview of t ...

classifies this family in the

suborder Order () 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 recognized ...

Ceratioidei Ceratioidei, the deep-sea anglerfishes or pelagic anglerfishes, is a suborder of marine ray-finned fishes, one of five suborders in the order Lophiiformes, the anglerfishes. These fishes are found in tropical and temperate seas throughout the wor ...

of the

order Order, ORDER or Orders may refer to: * A socio-political or established or existing order, e.g. World order, Ancien Regime, Pax Britannica * Categorization, the process in which ideas and objects are recognized, differentiated, and understood ...

Lophiiformes The anglerfish are ray-finned fish in the order Lophiiformes (). Both the order's common and scientific name comes from the characteristic mode of predation, in which a modified dorsal fin ray acts as a lure for prey (akin to a human angler, ...

Species and
species groups In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...

There are currently 23 recognized species in this genus and these are divided into species groups as set out below. * ''groenlandicus'' group ** '' Himantolophus crinitus'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus danae'' Regan & Trewavas, 1932 ** '' Himantolophus groenlandicus'' J. C. H. Reinhardt, 1837 (Atlantic footballfish) ** '' Himantolophus paucifilosus'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus sagamius'' S. Tanaka (I), 1918 (Pacific footballfish) * ''appelii'' group ** '' Himantolophus appelii'' F. E. Clarke, 1878 (Prickly anglerfish) ** '' Himantolophus stewarti'' Pietsch & Kenaley, 2011 * ''nigricornis'' group ** '' Himantolophus melanolophus'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus nigricornis'' Bertelsen & G. Krefft, 1988 *''albinares'' group ** ''

Himantolophus albinares ''Himantolophus albinares'' is a species of footballfish, a type of anglerfish. The fish is bathypelagic and can be found at depths ranging from . It is endemic to the Atlantic Ocean The Atlantic Ocean is the second largest of the world's ...

Maul A maul may refer to any number of large hammers, including: * War hammer, a medieval weapon * Post maul, a type of sledgehammer * Spike maul, railroad hand tool * Splitting maul, heavy wood-splitting tool resembling both axe and hammer People * A ...

, 1961 ** '' Himantolophus borealis'' Kharin, 1984 ** ''

Himantolophus kalami ''Himantolophus kalami'' is a species of marine Actinopterygii, ray-finned fish belonging to the Family (biology), family Himantolophidae, the footballfishes. This species was Species description, described in 2022 by Melapurra Rajeeshkumar, Theod ...

'' Rajeeshjumar, Pietsch & Saravanane, 2022 ** '' Himantolophus mauli'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus multifurcatus'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus pseudalbinares'' Bertelsen & G. Krefft, 1988 * ''cornifer'' group ** '' Himantolophus azurlucens'' Beebe & Crane, 1947 ** '' Himantolophus compressus'' Osório, 1912 ** '' Himantolophus cornifer'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus litoceras'' A. L. Stewart & Pietsch, 2010 ** '' Himantolophus macroceras'' Bertelsen & G. Krefft, 1988 ** '' Himantolophus macroceratoides'' Bertelsen & G. Krefft, 1988 *''brevirostris'' group ** '' Himantolophus brevirostris'' Regan, 1925 * ''rostratus'' group ** '' Himantolophus rostratus'' Regan, 1925 These groups were determined from the morphology of the metamorphosed females, except for ''brevirostris'' and ''rostratus'' which were determined from males only.

Characteristics

Footballfish are sexually dimorphic with the metamorphosed females and males being very different in appearance. The metamorphosed females are distinguished from other anglerfishes of the suborder Ceratioidei by having a well developed lower jaw which protrudes beyond the snout. They also have a wide

vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...

which has no teeth, well developed spines on the sphenotic bone, there is a covering of low, rounded

papillae Papilla (Latin, 'nipple') or papillae may refer to: In animals * Papilla (fish anatomy), in the mouth of fish * Papilla (worms), small bumps on the surface of certain worms * Basilar papilla, a sensory organ of lizards, amphibians and fish * De ...

on the snout and chin and, at least in larger individuals, there are conical spines in the skin which are scattered over the head and body. The

esca Esca can refer to: * The esca or illicium, a modified luminescent fin ray found in anglerfish * Esca (grape disease) * the Ezka, a river in the Basque country known as "Esca" in Spanish * ESCA or Electron Spectroscopy for Chemical Analysis, now ...

of footballfishes vary in size and morphology, to a greater extent than other deep sea anglerfishes. The metamorphosed males have a line of large spines above and behind the upper denticular bone, their eyes are directed to the sides and have moderately sized and they have a large olfactory system with sideways pointing nostrils. They have between 16 and 31 denticular teeth on the snout and between 20 and 50 on the chin, these teeth merge at their bases to form the upper and lower denticular bones. Their skin has a dense covering of dermal

spinules Spinules are small spines or thorns (vertebral columns) that are part of biological and manmade structures. The word originates from the Latin word and is often used in botany and zoology. The presence or absence of spinules, and their shape, can ...

. The larvae are round with a swollen appearance to the skin with pectoral fins that do not extend beyond the dorsal and anal fins, the females have a small club-shaped rudimentary illicium. The males are considerably smaller than the females, for example in ''H. groenlandicus'' the maximum published

standard length Fish measurement is the measuring of individual fish and various parts of fish anatomy, their anatomies, for data used in many areas of ichthyology, including Taxonomy (biology), taxonomy and fishery biology. Overall length Standard length (SL) is ...

for a male is while that of a females is .

Distribution and habitat

Footballfishes are found in the

mesopelagic The mesopelagic zone (Greek language, 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

bathypelagic The bathypelagic zone or bathyal zone (from Greek βαθύς (bathýs), deep) is the part of the open ocean that extends from a depth of below the ocean surface. It lies between the mesopelagic above and the abyssopelagic below. The bathypela ...

zones in the Atlantic, Pacific and Indian Oceans, as well as the Southern Ocean.

Biology

Footballfishes are one of the ceratioid groups in which the males are free living and non-parasitic on the females. Despite their small size relative to the females, the males are the largest among the free-living male Ceratioids. The males use their highly developed olfactory organs to detect females, once they find a female they attach themselves to her but do not fuse with her (as in the parasitic species). The eggs and larvae are

pelagic The pelagic zone consists of the water column of the open ocean and can be further divided into regions by depth. The word ''pelagic'' is derived . The pelagic zone can be thought of as an imaginary cylinder or water column between the sur ...

. The specialised teeth on the denticular bones are used to temporarily attach the male to the female. There are records of 5 females with white, circular scars on their skin, thought to be caused by males that became detached somehow, but have alternatively been suggested to be damage from parasitic

copepod Copepods (; meaning 'oar-feet') are a group of small crustaceans found in nearly every freshwater and saltwater habitat (ecology), habitat. Some species are planktonic (living in the water column), some are benthos, benthic (living on the sedimen ...

s. At the depths at which these fishes live it is dark and food is sparse and rarely encountered. The female footballfish have

bioluminescent Bioluminescence is the emission of light during a chemiluminescence reaction by living organisms. Bioluminescence occurs in multifarious organisms ranging from marine vertebrates and invertebrates, as well as in some fungi, microorganisms inc ...

bacteria in their escas and this is used to attract prey to within striking distance of the mouth. The prey is whatever they can fit into their mouths, and the backward curving teeth ensure that prey are unable to escape. Recorded prey includes fishes, squid and crustaceans.

Gallery

File:Himantolophus groenlandicus by OpenCage.jpg File:Hal - Himantolophus groenlandicus 2.jpg File:Faroese stamp 539 atlantic footballfish.jpg

References

*Anderson, M. Eric and Leslie, Robin W. (2001)
"Review of the deep-sea anglerfishes (Lophiiformes: Ceratioidei) of southern Africa"
''Ichthyological Bulletin''. J.L.B. Smith Institute of Ichthyology, Rhodes University. . Retrieved October 31, 2005. *Munk, Ole. (1999)
"The escal photophore of ceratioids (Pisces; Ceratioidei) — a review of structure and function"
''

Acta Zoologica ''Acta Zoologica'' is a peer-reviewed scientific journal published by Wiley-Blackwell on behalf of the Royal Swedish Academy of Sciences and the Royal Danish Academy of Sciences and Letters. It is one of the world's leading zoological journals an ...

'', Vol 80., Issue 4, pp. 265–284. Retrieved October 31, 2005. {{Taxonbar, from1=Q9292308, from2=Q541816 Himantolophidae Ceratioidei Deep sea fish Marine fish genera Ray-finned fish genera