Beloniformes

(vertebrate zoology) The former ordinal name for a group of fishes now included in the order Atheriniformes.

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(Needlefishes and relatives)

Class: Actinopterygii

Order: Beloniformes

Number of families: 5

Evolution and systematics

The Beloniformes is one of three orders within the series Atherinomorpha. One of the other two orders, the Atheriniformes, is thought by some ichthyologists to represent an unnatural grouping of several lineages, while others consider it monophyletic (a natural group). The other order, Cyprinodontiformes, is the sister-group of the Beloniformes, as evidenced by numerous internal characters, including modifications to the gill arches and the bones surrounding the eyes. Both Cyprinodontiformes and Beloniformes are agreed to be monophyletic. Beloniformes themselves are united by derived internal characters of the gill arches rather than any conspicuous external morphological characters. Five families of fishes make up the order Beloniformes: Adrianichthyidae (ricefishes), Belonidae (needlefishes), Scomberesocidae (sauries), Exocoetidae (flyingfishes), and Hemiramphidae (halfbeaks). Within these families are 38 genera and about 200 species, 51 of which are either brackish or freshwater, the remainder of which are marine.

The earliest known fossil Beloniformes are just over 50 million years old, and come from two sites: the exocoetids from Monte Bolca in Italy, and the hemiramphids from the Selsey formation in England. Beloniformes are broadly divided into two suborders, the Exocoetoidei (beaked forms: belonids, scomberesocids, exocoetids, and hemiramphids) and the Adrianichthyoidei (lacking a beak; adrianichthyids). Adrianichthyids were traditionally included within the Cyprinodontiformes until Rosen and Parenti argued for their inclusion within the Beloniformes in 1981, based mainly on characters of the gill arches and hyoid apparatus. Adrianichthyids are now considered to be the sister lineage to the rest of the order, within which the sister groups Belonidae-Scomberesocidae and Hemiramphidae-Exocoetidae have been suggested.

Contrary to this scheme, a study based on morphology and molecules, published in 2000 by Lovejoy, places halfbeaks as the ancestral form among the beaked beloniforms. In Love-joy's tree, some halfbeaks are most closely related to needle-fishes and sauries, while the marine halfbeak Hemiramphus is sister to flyingfishes. This result refutes an old hypothesis, based on the observation that needlefishes pass through a half-beak stage during their development, that halfbeaks derive from needlefishes via truncation of the development sequence.

Physical characteristics

Beloniformes are typically elongate fishes, with dorsal and anal fins situated posteriorly on the body and the lateral line situated ventrally. Additional characteristics of the group include fusion of the toothed 5th ceratobranchials into a lower pharyngeal jaw and an open nasal pit. Belonids, aptly called needlefishes, are sleek and garlike piscivores, with very long upper and lower jaws studded with sharp teeth. They can achieve lengths of up to 3.3 ft (1 m). A small number of needle-fishes have a reduced upper jaw, and like halfbeaks, feed on plankton and insects. Scomberesocids, of which the largest are about 1.65 ft (0.5 m) long, can be distinguished from belonids by the five or six finlets behind their dorsal and anal fins. The diminutive scomberesocid Cololabis adocetus, at 3 in (7.5 cm), is the smallest fish in the surface waters of the open ocean.

In most species of hemiramphids, or halfbeaks, the lower jaw is much longer than the upper. The front margin of the upper jaw is triangular in shape, the scales are large and cycloid, and fin spines are lacking. Exocoetids, the flyingfishes, are torpedo shaped with greatly enlarged pectoral fins, and the lower lobe of the caudal fin is stiffened and much larger than the upper. Interestingly, the most primitive flyingfish genera, Oxyporhamphus (once included with hemiramphids),

Fodiator, and Parexocoetus, have elongate lower jaws reminiscent of halfbeaks. More derived flyingfishes have acquired oversized pelvic fins in addition to large pectoral fins, and are called four-wingers.

Adrianichthyids, the most basal among the Beloniformes, are superficially unlike other members of the group. Most of the species are in the genus Oryzias, and are small, relatively deep-bodied fishes with large eyes, upturned mouths, and a long anal fin base. Noteworthy in the family is the duckbilled buntingi (Adrianichthys kruyti), which has a bill-shaped mouth with the upper jaw overhanging the lower. Xenopoecilus species also have a bill-shaped mouth, and the carry their eggs at the base of the pelvic fins by way of filamentous attachments.

Distribution

Beloniformes are widely distributed in temperate and tropical marine and fresh waters. Adrianichthyids are found in Asian fresh and brackish waters from India to Japan, and in the Indo-Australian archipelago. Belonids are found in the open ocean in tropical and temperate seas worldwide, with numerous species in the freshwaters of South America and some in Asia. Like marine belonids, scomberesocids are widely distributed in warmer waters of the open ocean. Exocoetids are found in warm waters of the Atlantic, Pacific, and Indian Oceans. Hemiramphids have a similar distribution in marine waters, but have also invaded freshwaters, especially in the Indo-Australian region.

Habitat

Marine beloniforms can be found in the surface waters of the open ocean, as well as in coastal habitats like estuaries and mangrove swamps. Adrianichthyids, belonids, and hemiramphids can be found in a diversity of tropical freshwater habitats, including lakes and rivers.

Behavior

One characteristic of many beloniforms is a strong attraction to lights at night. This behavior is exploited by fishermen, who use lights to capture schools of sauries that cruise just below the ocean surface. Such fishing methods, however, involve a peculiar (although infrequent) hazard: impalement by large needlefishes. In one documented case, a 3.3 ft (1 m) long Tylosurus fatally impaled a fisherman after jumping toward the light on board his canoe.

Certainly the most remarkable beloniform behavior is exocoetid flight. (It should be noted that flight is not entirely restricted to exocoetids: Some hemirhamphids are capable of gliding, and Euleptorhamphus viridis has been reported to travel 164 ft [50 m] in two jumps.) In the more derived four-winged flyingfish species, flight is achieved as follows. The fish, swimming at a speed of about 33 ft/s (10 m/s), breaks the surface at an oblique angle and taxis for 16.4–82 ft (5–25 m) by rapidly beating the caudal fin in the water. Then a free flight ensues, which may span a distance of 164 ft (50 m) and reach a height of 26.2 ft (8 m). Once the fish loses altitude, caudal fin taxiing can be repeated without returning to the water, so that flights can be stretched to distances of 1,312 ft (400 m). Intriguingly, flyingfishes seem to sense wind direction and take off into the wind, and tantalizing evidence suggests that they can control the direction of flight; Cypsilurus appear to successfully seek out patches of seaweed in which to land. So why do flyingfishes fly? One of the most likely hypotheses is that flight has evolved as a tactic for evading predators.

Feeding ecology and diet

Beloniformes utilize a relatively wide spectrum of foods. Most impressive perhaps are the marine needlefishes, which cruise through the surface waters of the open ocean devouring small fishes. However, not all needlefishes are piscivores. In the Amazon, many belonids feed heavily on zooplankton or insects. Belonion apodion, which grows only to about 2 in (5 cm), is unusual in that it deftly snaps up individual rotifers, which are less than 0.004 in (0.1 mm) long and usually pass through the gill rakers of filter-feeding planktivorous fishes. Potamorrhaphis, which prefers terrestrial insects (especially flying ants), hovers motionlessly and waits for prey to fall to the surface alongside its body. Then it rapidly curls the body and strikes at the prey from the side.

Freshwater halfbeaks also feed on terrestrial insects, and some are particularly well suited to this mode of feeding. Members of the genus Hemirhamphodon are noteworthy for having numerous anteriorly directed teeth on their lower jaws, which ensnare ants and other insects found floating on the surface. Marine halfbeaks, on the other hand, tend to feed on algae, diatoms, and sea grasses, though some species eat small fishes. Planktivorous marine beloniforms include the flyingfishes and sauries. Ricefishes are omnivorous and will eat plankton, small insects, detritus, and plant material.

Beloniformes themselves often fall prey to larger fishes. Flyingfishes in particular are eaten by mackerel, tuna, and marlin, among other predatory fishes, as well as squids and birds.

Reproductive biology

Much of what is known about beloniform reproductive biology involves the eggs and larvae. Typically, eggs develop in one to two weeks, and the larvae are immediately able to feed upon hatching. Many pelagic beloniform eggs have filamentous projections that cause them to stick to floating debris. Needlefish eggs have tendrils that are particularly sticky, and they form egg clusters that stick to other objects in the water. Likewise, sauries produce filamentous eggs that float in open water, but they are less adhesive than needlefish eggs. Flyingfishes lay pelagic eggs that may or may not have filaments, and some species attach their eggs to floating seaweed. Marine halfbeaks lay eggs with tendrils that float about in open water, but some freshwater representatives bear live young, namely Dermogenys, Nomorhamphus, and Hemirhamphodon. In these viviparous forms, long genital papillae are used for internal fertilization, and the male anal fin is modified into an andropodium.

The adrianichthyid Horaichthys from India, uniquely among atherinomorphs, produces encapsulated sperm bundles, or spermatophores. In adrianichthyids other than Oryzias, fertilization is apparently internal. The eggs of many species of adrianichthyids are retained externally by the female for various lengths of time. Females of the species Xenopoecilus oophorus, known as the egg-carrying buntingi, carry a cluster of about 30 fertilized eggs attached by filaments in an external concavity near the vent. The pelvic fins cover and protect this egg mass.

Conservation status

No Beloniformes are CITES listed, or listed as endangered by the U. S. Fish and Wildlife Service. However, 16 species are included on the IUCN Red List. Eight of those, mostly Oryzias species, are categorized as Vulnerable, one species is listed as Lower Risk/Near Threatened, and two species are listed as Data Deficient. Three species are listed as Endangered: Oryzias orthognathus, Xenopoecilus oophorus, and Xenopoecilus sarasinorum. Listed as Critically Endangered are Adrianichthys kruyti and Xenopoecilus poptae, both of which are known only from Lake Poso, Sulawesi. Although no Beloniformes are formally listed as Extinct in the Wild by the U. S. Fish and Wildlife Service or Extinct by the IUCN, Adrianichthys kruyti is generally thought to be extinct. Pressure from an introduced species of catfish, in addition to parasites that entered the lake with the catfish, are implicated in the decline of the Lake Poso adrianichthyids.

Significance to humans

Many Beloniformes are fished at night using lights, and some rather creative methods have been devised. Where flyingfishes are abundant, fishermen that leave a light suspended all night over a canoe partially filled with water can return in the morning to a boat full of fresh fish. The fish are drawn to the light and jump into the canoe, but have too little water to jump back out. Flyingfishes are also attracted to leaves or straw scattered about the surface as a place to lay their eggs, and can be fished by using such material.

Some Beloniformes are used by humans as more than just food. Numerous freshwater species, including halfbeaks, ricefishes, and needlefishes, can be found in the aquarium trade. In Thailand, the halfbeak, Dermogenys pusillus, is bred in captivity so that males, which will engage rivals by locking jaws, can be used as fighting fish. Members of the genus Oryzias are propagated in large numbers in captivity to be used in experimental research.

Finally, it should be mentioned that needlefishes can in rare cases be traumatogenic, causing injury or death by means of impalement. In one such case, a hapless surfer was killed when the snout of a fast-swimming needlefish went through his eye socket and into his brain.

Species accounts

Resources

Berra, T. M. Freshwater Fish Distribution. San Diego: Academic Press, 2001.

Breder, C. M., Jr., and D. E. Rosen. Modes of Reproduction in Fishes. Garden City, NY: The Natural History Press, 1966.

Collette, B. B., G. E. McGowen, N.V. Parin, and S. Mito. "Beloniformes: Development and Relationships." In Ontogeny and Systematics of Fishes, edited by H. G. Moser. Lawrence, KS: Allen Press, 1984.

Fischer, W., F. Krupp, W. Schneider, C. Sommer, K. E. Carpenter, and V. Niem, eds. Guia FAO para Identification de Especies para lo Fines de la Pesca. Pacifico Centro-Oriental, Vol. 2. Rome: FAO, 1995.

Harrison, I. J., and M. L. J. Stiassny. "The Quiet Crisis: A Preliminary Listing of the Freshwater Fishes of the World that Are Extinct or 'Missing in Action.'" In Extinctions in Near Time, edited by R. D. E. MacPhee. New York: Kluwer Academic/Plenum Publishers, 1999.

Nelson, J. S. Fishes of the World. 3rd edition. New York: John Wiley & Sons, 1994.

Parin, N. V. "Exocoetidae: Flyingfishes." In FAO Species Identification Guide for Fishery Purposes. The Living Marine Resources of the Western Central Pacific, Vol. 4. Bony Fishes, Part 2 (Mugilidae to Carangidae), edited by K. E. Carpenter and V. H. Niem. Rome: FAO, 1999.

Patterson, C. "Osteichthyes: Teleostei." In The Fossil Record 2, edited by M. J. Benton. London: Chapman and Hall, 1993.

Paxton, J. R., and W. N. Eschmeyer, eds. Encyclopedia of Fishes. 2nd edition. San Diego: Academic Press, 1998.

Riehl, R., and H. A. Baensch. Aquarium Atlas. Melle, Germany: Baensch, 1986.

Anderson, W. D., III, and B. B. Collette. "Revision of the Freshwater Viviparous Halfbeaks of the Genus Hemirhamphodon (Teleostei: Hemiramphidae)." Ichthyological Exploration of Freshwaters 2 (1991): 151–176.

Boughton, D. A., B. B. Collette, and A. R. McCune. "Heterochrony in Jaw Morphology of Needlefishes (Teleostei: Belonidae)." Systematic Zoology 40 (1991): 329–354.

Coates, D., and P. A. M. Van-Zwieten. "Biology of the Freshwater Halfbeak Zenarchopterus kampeni (Teleostei: Hemiramphidae) from the Sepik and Ramu River Basin, Northern Papua New Guinea." Ichthyological Exploration of Freshwaters 3 (1992): 25–36.

Dasilao, J. C., Jr., and K. Sasaki. "Phylogeny of the Flyingfish Family Exocoetidae (Teleostei, Beloniformes)." Ichthyological Research 45 (1998): 347–353.

Davenport, J. "How and Why do Flying Fish Fly?" Reviews in Fish Biology and Fisheries 4 (1994): 184–214.

Goulding, M., and M. L. Carvalho. "Ecology of Amazonian Needlefishes (Belonidae)." Revista Brasileira de Zoologia 2(1984): 99–111.

Kottelat, M. "Synopsis of the Endangered Buntingi (Osteichthyes: Adrianichthyidae and Oryziidae) of Lake Poso, Central Sulawesi, Indonesia, with a New Reproductive Guild and Descriptions of Three New Species." Ichthyological Exploration of Freshwaters 1 (1990): 49–67.

Lovejoy, N. R. "Reinterpreting Recapitulation: Systematics of Needlefishes and Their Allies (Teleostei: Beloniformes)." Evolution 54 (2000): 1,349–1,362.

Lovejoy, N. R., and B. B. Collette. "Phylogenetic Relationships of New World Needlefishes (Teleostei: Belonidae) and the Niogeography of Transitions Between Marine and Freshwater Habitats." Copeia 2001, no. 1(2001): 324–338.

Lovejoy, N. R., and M. L. G. De-Araujo. "Molecular Systematics, Biogeography and Population Structure of Neotropical Freshwater Needlefishes of the Genus Potamorrhaphis." Molecular Ecology 9 (2000): 259–268.

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Mok, E. Y. M., and A. D. Munro. "Observations on the Food and Feeding Adaptations of Four Species of Small Pelagic Teleosts in Streams of the Sungei Buloh Mangal, Singapore." Raffles Bulletin of Zoology 39 (1991): 235–257.

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[Article by: Robert Schelly, MA]

Beloniforms
Exocoetus volitans
Scientific classification
Kingdom: Animalia Phylum: Chordata Class: Actinopterygii Order: Beloniformes
Families
Suborder Adrianichthyoidei Adrianichthyidae Suborder Belonoidei Superfamily Scomberesocoidea Belonidae Scomberesocidae Superfamily Exocoetoidea Exocoetidae Hemiramphidae

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Beloniformes

The Beloniformes are an order of five families of freshwater and marine ray-finned fish: the Adrianichthyidae (ricefish and medakas): Belonidae (needlefish); Exocoetidae (flyingfishes); Hemiramphidae (halfbeaks): and the Scomberesocidae (sauries).^[1] With the exception of the Adrianichthyidae, these are streamlined, medium-sized fishes that live close to the surface of the water feeding on algae, plankton, or smaller animals including other fishes. Most are marine, though a few needlefish and halfbeaks inhabit brackish and fresh waters.^[2]

The order is sometimes divided up into two suborders, the Adrianichthyoidei and the Belonoidei. The Adrianichthyoidei contains only a single family, the Adrianichthyidae. Originally, the Adrianichthyidae were included in the Cyprinidontiformes and assumed to be closely related to the killifish, but a closer relationship to the beloniforms is indicated by various characteristics including the absence of the interhyal, resulting in the upper jaw being fixed or nonprotrusible. The Belonoidei may also be further subdivided into two superfamilies, the Scomberesocoidea and the Exocoetoidea. The Scomberesocoidea contains the Belonidae and Scomberesocidae, while the Exocoetoidea comprises the Exocoetidae and Hemiramphidae.^[3] However, newer evidence finds that the flyingfishes are nested within the halfbeaks, and the needlefish and sauries are nested within the subfamily Zenarchopterinae of the family Hemiramphidae, which has been recognized as its own family. The sauries are also nested within the family Belonidae.^[4]

The beloniforms display an interesting array of jaw morphologies. The basal condition in the order excluding the ricefishes is an elongated lower jaw in juveniles and adults as represented in halfbeaks. In the needlefish and sauries, both jaws are elongated in the adults; the juveniles of most species develop through a "halfbeak stage" before having both jaws elongated. The elongated lower jaw is lost in adults and is lost in most juveniles in the flyingfishes and some halfbeak genera.^[4]

See also

• Gallery of beloniform fishes

References

1. ^ Froese, R. and D. Pauly. Editors.. Order Summary for Beloniformes. FishBase. Retrieved on 2007-02-10.
2. ^ Helfman G., Collette B., & Facey D.: The Diversity of Fishes, Blackwell Publishing, pp 274-276, 1997, ISBN 0-86542-256-7
3. ^ Nelson, Joseph S. (2006). Fishes of the World. John Wiley & Sons, Inc. ISBN 0-471-25031-7. 
4. ^ ^{a b} Lovejoy, N; Iranpour, M; Collette, B (2004). "Phylogeny and Jaw Ontogeny of Beloniform Fishes". Integrative and Comparative Biology 44: 366–377. 

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