Anglerfish

, Le Règne Animal. Toadfish (Batrachoididae) such as Batrachoides surinamensis (middle) are no longer considered close to anglerfish Anglerfish were first grouped in the family of Acanthopterygians with "pediculate pectoral [fin]s" () by Cuvier in the 1829 edition of Le Règne Animal; being characterized by possessing "a sort of arm supporting their pectorals, formed by an elongated carpal bone". Cuvier placed the genera Lophius (incl. Lophius piscatorius), Chironectes/Antennarius (incl. various subspecies of Lophius histrio), Malthe (incl. Lophius vespertilio), and Batrachus within this family. which was eventually truncated into Pediculati or Pediculata (pediculate fish), these names being used to classify anglerfish through 1926. Though this term saw use in publications as late as the 1970s, Pediculati has fallen out of use. The group Lophidia was conceived by Samuel Garman in 1899; this group was subdivided into the Lophioids (incl. Lophius, Lophiomus, Melanocetus, Dolopichthys, Chaunax, and Chaunacops) and the Halieutoids (incl. Oncocephalus, Halieutaea, Halieutella, Halieutichthys, Halieutopsis, Halicmetus, Dibranchus, Dibranchichthys, and Malthopsis) based on the orientation of the ilicium's base. Classification The following classification is based on ''Eschmeyer's Catalog of Fishes'' (2025): • Suborder Lophioidei Regan, 1912 • Family Lophiidae Rafinesque, 1810 (monkfishes and goosefishes) • Suborder Ogcocephaloidei Pietsch, 1984 • Family Ogcocephalidae Gill, 1893 (batfishes) • Suborder Antennarioidei Regan, 1912 • Family Antennariidae Jarocki, 1822 (frogfishes) • Subfamily Fowlerichthyinae Maile, Smith & Davis, 2025 (fanfin frogfishes) • Subfamily Antennariinae Jarocki, 1822 (Fibonacci frogfishes) • Subfamily Lophichthyinae Boeseman, 1964 (lophichthyin frogfishes) • Subfamily Tathicarpinae Hart, Arnold, Alda, Kenaley, Pietsch, Hutchinson & Chakrabarty, 2022 (longfin frogfishes) • Subfamily Tetrabrachiinae Regan, 1912 (tetrabrachiid frogfishes) • Subfamily Histiophryninae Arnold & Pietsch, 2012 (starfingered frogfishes) • Subfamily Rhycherinae Hart, Arnold, Alda, Kenaley, Pietsch, Hutchinson & Chakrabarty, 2022 (Balrog frogfishes) • Subfamily Brachionichthyinae Gill, 1863 (handfishes) • Suborder Chaunacoidei Pietsch & Grobecker, 1987 • Family Chaunacidae Gill, 1863 (gapers or sea toads) • Suborder Ceratioidei Regan, 1912 • Family Caulophrynidae Goode & Bean, 1896 (fanfins) • Family Neoceratiidae Regan, 1926 (spiny seadevils) • Family Melanocetidae Gill, 1878 (black seadevils) • Family Himantolophidae Gill, 1861 (footballfishes) • Family Diceratiidae Regan & Trewavas, 1932 (double anglers) • Family Oneirodidae Gill, 1878 (dreamers) • Family Thaumatichthyidae Smith & Radcliffe, 1912 (wolftrap anglers) • Family Centrophrynidae Bertelsen, 1951 (prickly seadevils) • Family Ceratiidae Gill, 1861 (warty seadevils) • Family Gigantactinidae Boulenger, 1904 (whipnose anglers) • Family Linophrynidae Regan, 1925 (leftvents) Alternatively, Lophiiformes may be treated as clade within Acanthuriformes; a 2025 paper defines Lophioidei as equivalent to the prior conception of Lophiiformes (the one depicted above) and converts the suborders into infraorders (as seen below). Phylogenetic studies have consistently recovered the Lophiiformes as sister-group to the Tetraodontiformes, The Lophiiformes and Tetraodontiformes are united by several derived morphological features separating them from other Acanthuriformes, including restricted gill openings, along with the absence of multiple skeletal elements, such as spines supporting the anal fin, ribs, nasals, and basisphenoid. == Evolution ==

The earliest fossils of anglerfish are from the Eocene, excavated from the Monte Bolca formation of Italy, and these already show evidence of diversification into the modern families that make up the order. Given this, and their close relationship to the Tetraodontiformes which are known from Cretaceous fossils, they likely originated during the Cretaceous. A 2010 mitochondrial genome phylogenetic study suggested the anglerfishes diversified in a short period during the early to mid-Cretaceous, between 130 and 100 million years ago. A 2024 study found that all anglerfish suborders most likely diverged from one another during the Late Cretaceous and Paleocene, but the multiple families of deep-sea anglerfishes (Ceratioidei), as well as their trademark sexual parasitism, originated during the Eocene in a rapid radiation following the Paleocene-Eocene thermal maximum. Adaptations to different ranges of depths may have driven the evolution of anglerfish species and families in prehistory. ImageSize = width:1200 height:auto barincrement:15px PlotArea = left:10px bottom:50px top:10px right:10px Period = from:-59 till:10 TimeAxis = orientation:horizontal ScaleMajor = unit:year increment:5 start:-59 ScaleMinor = unit:year increment:1 start:-59 TimeAxis = orientation:hor AlignBars = justify Colors = #legends id:CAR value:claret id:ANK value:rgb(0.4,0.3,0.196) id:HER value:teal id:HAD value:green id:OMN value:blue id:black value:black id:white value:white id:cenozoic value:rgb(0.54,0.54,0.258) id:paleogene value:rgb(0.99,0.6,0.32) id:paleocene value:rgb(0.99,0.65,0.37) id:eocene value:rgb(0.99,0.71,0.42) id:oligocene value:rgb(0.99,0.75,0.48) id:neogene value:rgb(0.999999,0.95,0.68) id:miocene value:rgb(0.999999,0.9,0.1) id:pliocene value:rgb(0.999999,0.999999,0.24) id:quaternary value:rgb(0.999999,0.999999,0.24) id:pleistocene value:rgb(0.97,0.98,0.68) id:holocene value:rgb(0.999,0.95,0.88) BarData= bar:NAM1 bar:NAM2 bar:NAM3 bar:NAM4 bar:NAM5 bar:NAM6 bar:NAM7 bar:NAM8 bar:NAM9 bar:NAM10 bar:NAM11 bar:NAM12 bar:space bar:period bar:space bar:era PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 shift:(7,-4) PlotData= align:left fontsize:M mark:(line,white) width:5 anchor:till align:left color:eocene bar:NAM1 from: -55.8 till: 0 text: Lophius color:eocene bar:NAM2 from: -55.8 till: 0 text: Brachionichthys color:eocene bar:NAM3 from: -55.8 till: 0 text: Antennarius color:eocene bar:NAM4 from: -48.6 till: 0 text: Ogcocephalus color:eocene bar:NAM5 from: -37.2 till: 0 text: Dibranchus color:eocene bar:NAM6 from: -37.2 till: 0 text: Chaunax color:miocene bar:NAM7 from: -11.63 till: 0 text: Oneirodes color:miocene bar:NAM8 from: -8.6 till: 0 text: Borophryne color:miocene bar:NAM9 from: -8.6 till: 0 text: Chaenophryne color:miocene bar:NAM10 from: -8.6 till: 0 text: Leptacanthichthys color:miocene bar:NAM11 from: -8.6 till: 0 text: Linophryne color:miocene bar:NAM12 from: -8.6 till: 0 text: Acentrophryne PlotData= align:center textcolor:black fontsize:M mark:(line,black) width:25 bar:period from: -59.0 till: -55.8 color:paleocene text:Paleocene from: -55.8 till: -33.9 color:eocene text:Eocene from: -33.9 till: -23.03 color:oligocene text:Oligocene from: -23.03 till: -5.332 color:miocene text:Miocene from: -5.332 till: -2.588 color:pliocene text:Plio. from: -2.588 till: -0.0117 color:pleistocene text:Pleist. from: -0.0117 till: 0 color:holocene text:H. bar:era from: -59.0 till: -23.03 color:paleogene text:Paleogene from: -23.03 till: -2.588 color:neogene text:Neogene from: -2.588 till: 0 color:quaternary text:Q. TextData = pos:(600,20) text:"Time (in million years)" == Anatomy ==

'', up to TL) is a frogfish well-adapted to live among sargassum Anglerfish are defined by gills that open behind the pectoral fins (as opposed to other fish whose pectorals lay behind the gill opening), depressible teeth that can hinge back, joints of the epiotic bone, the form of the pectoral fin radials, and the luring apparatus (see subsection). The largest members are the European monkfish Lophius piscatorius ( SL, ), the deep-sea warty anglerfish Ceratias holboelli ( TL), the giant frogfish Antennarius commerson ( TL), and the giant triangular batfish Malthopsis gigas (). Many suborders are sexually dimorphic, with the deep-sea anglerfish being the most extreme example; male C. holboelli can reach up to long (SL), while females are commonly around TL, Sexual dimorphism is not as pronounced in other suborders; the Lophiid monkfish genus Lophiodes are quite similar in size between the genders (Mean for Males SL; Females SL), and the same is true for Lophius itself (Males ; Females ). To blend in with the featureless dark depths they inhabit, deep-sea anglerfish are dark colored, with tints ranging from grey to brown. Anglerfish are able to distend both their jaw and stomach to enormous size, since their bones are thin and flexible, which allows them to swallow prey up to twice as large as their entire bodies. The illicial apparatus is most notable in the deep-sea anglerfish (Ceratioidei) as their esca contain bioluminescent bacteria, making them glow in the dark waters of the deeper pelagic zones. In at least the triplewart seadevil, the illicium is moved back and forth by five distinct pairs of muscles: namely the shorter erector and depressor muscles that dictate movement of the illicial bone, along with inclinator, protractor, and retractor muscles that aid motion of the pterygiophore. == Behavior ==

Predation '': The first spine of the dorsal fin of the anglerfish acts like a fishing rod with a lure. The name "anglerfish" derives from the species' characteristic method of predation. Anglerfish typically have at least one long filament sprouting from the middle of their heads, termed the illicium. The illicium is the detached and modified first three spines of the anterior dorsal fin. In most anglerfish species, the longest filament is the first. This first spine protrudes above the fish's eyes and terminates in an irregular growth of flesh (the esca), and can move in all directions. Anglerfish can wiggle the esca to make it resemble a prey animal, which lures the anglerfish's prey close enough for the anglerfish to devour them whole. Some deep-sea anglerfish of the bathypelagic zone also emit light from their esca to attract prey. Because anglerfish are opportunistic foragers, they show a range of preferred prey with fish at the extremes of the size spectrum, whilst showing increased selectivity for certain prey. One study examining the stomach contents of threadfin anglerfish (Lophiodes spilurus) off the Pacific coast of Central America found these fish primarily ate two categories of benthic prey: crustaceans and teleost fish. The most frequent prey were pandalid shrimp. 52% of the stomachs examined were empty, supporting the observations that anglerfish are low energy consumers. Movement and energy conservation "standing" on the benthos All anglerfish are weak swimmers, including the pelagic deep-sea anglerfish. Demersal species often "walk" on the bottom upon their pectoral and pelvic fins. The pelvic fins were lost in the deep-sea anglers. The jaw and stomach of the anglerfish can extend to allow it to consume prey up to twice its size. Because of the limited amount of food available in the anglerfish's environment, this adaptation allows the anglerfish to store food when there is an abundance. The sea toad Chaunax endeavouri has been observed to retain water in its gills for at least around 26 seconds and up to 4 minutes in some cases. This behavior is thought to be an energy-saving measure as respiration requires energy, thus the fish "holding its breath" may conserve enough energy for such a behavior to be beneficial. Reproduction : Haplophryne mollis female anglerfish with males attached The deep-sea anglerfish employ an unusual mating method: because individuals are locally rare, encounters between two of the same species are also very rare, and finding a mate can be problematic; this has led to the development of sexual parasitism in anglerfish, where the males latch onto their mates using their mouths, which may not be suitable or effective for prey capture. When scientists first started capturing ceratioid anglerfish, they noticed that all of the specimens were female, and on some of these they had what appeared to be parasites attached to them, which turned out to be highly dimorphic male ceratioids. This is one of the few instances of naturally occurring parabiosis. The spawn of all anglerfish are enveloped by a gelatinous sheath, which has multiple terms referring to it. == Relation to humans ==

Classical interpretation In the History of Animals, Aristotle described the "Fishing-Frog" (one of the local Lophius species, like L. piscatorius or L. budegassa) as an example of a marine species well adapted to their environment, those equipped with "ingenious devices" that it uses to capture prey, alongside the Torpedo. He noted that fishing-frogs that have lost their lure appeared to be thinner than those still intact. In Asia, especially Japan, monkfish liver, known as ankimo, is considered a delicacy. Anglerfish is especially heavily consumed in South Korea, where it is featured as the main ingredient in dishes such as Agujjim. Northwest European Lophius species are heavily fished and are listed by the ICES as "outside safe biological limits". In 2010, Greenpeace International added the American angler (Lophius americanus), the angler (Lophius piscatorius), and the black-bellied angler (Lophius budegassa) to its seafood red list—a list of fish commonly sold worldwide with a high likelihood of being sourced from unsustainable fisheries. Additionally, anglerfish are known to occasionally rise to the surface during El Niño, leaving large groups of dead anglerfish floating on the surface. though these are all species that inhabit shallow waters; deep-sea anglerfish have not been kept in captivity due to the challenges of keeping them alive through capture, transport, and a display that can repressurize them. Antennarius biocellatus is known by the common names brackish-water frogfish or freshwater frogfish; being euryhaline, it can live in freshwater for some time, sometimes claimed to be the sole representative of the anglerfish to live in freshwater. Like many frogfish, it has been displayed in public aquaria, though unlike the other species A. biocellatus are sometimes kept in home aquaria by private aquarists. == References ==