Halfbeak

{{Short description|Family of fishes}} {{for|the United States Navy submarine|USS Halfbeak (SS-352)}} {{Automatic taxobox | fossil_range = {{Fossil range|50|0|Early Eocene to present}} | image = Halfbeak Cuba.JPG | image_caption = Ballyhoo (''Hemiramphus brasiliensis''), a marine halfbeak | taxon = Hemiramphidae | authority = T. N. Gill, 1859 | subdivision_ranks = Genera | subdivision = see text }}

'''Hemiramphidae''' is a family of fishes that are commonly called '''halfbeaks''', '''spipe fish''' or '''spipefish'''.{{Citation needed|date=May 2026|reason=Spipefish?}} They are a geographically widespread and numerically abundant family of epipelagic fish inhabiting warm waters around the world. The halfbeaks are named for their distinctive jaws, in which the lower jaws are significantly longer than the upper jaws. The similar viviparous halfbeaks (family Zenarchopteridae) have often been included in this family.

Though not commercially important themselves, these forage fish support artisanal fisheries and local markets worldwide. They are also fed upon by other commercially important predatory fishes, such as billfishes, mackerels, and sharks.

==Taxonomy== thumb|250px|left|''Oxyporhamphus micropterus'' has been considered either a halfbeak or a flyingfish. In 1758, Carl Linnaeus was the first to scientifically describe a halfbeak, ''Esox brasiliensis'' (now ''Hemiramphus brasiliensis''). In 1775 Peter Forsskål described two more species as ''Esox'', ''Esox far'' and ''Esox marginatus''. In 1816, Georges Cuvier created the genus ''Hemiramphus''; from then on, all three were classified as ''Hemiramphus''. In 1859, Theodore Nicholas Gill erected the Hemiramphidae, deriving their name from ''Hemiramphus'', the family's type genus.<ref name="Collette"/> The name comes from the Greek ''hemi'', meaning half, and ''rhamphos'', meaning beak or bill.<ref name= "fishbase">{{FishBase family|family= Hemiramphidae|year=2007|month= Aug}}</ref>

Currently, eight genera (including 60 species) are placed within the family Hemiramphidae:<ref name = CofF>{{Cof family|family= Hemiramphidae| access-date=18 August 2019}}</ref>

* ''Arrhamphus'' Günther, 1866 * ''Chriodorus'' Goode & Bean, 1882 * ''Euleptorhamphus'' Gill, 1859 * ''Hemiramphus'' Cuvier, 1816 * ''Hyporhamphus'' Gill, 1859 * ''Melapedalion'' Fowler, 1934 * ''Oxyporhamphus'' Gill, 1864 * ''Rhynchorhamphus'' Fowler, 1928

This family is primarily marine and found in the Atlantic, Pacific, and Indian Oceans, though some inhabit estuaries and rivers.

==Evolution== The halfbeaks' fossil record extends into the Lower Tertiary.<ref name="fishbase"/> The earliest known halfbeak is "Hemiramphus" ''edwardsi'' from the Eocene at Monte Bolca, Italy, although its attribution to the extant ''Hemiramphus'' is doubtful.<ref>{{Cite journal |last1=Bannikov |first1=A. F. |last2=Carnevale |first2=G. |last3=Kotlyar |first3=A. N. |date=2016-11-01 |title=A new halfbeak species (Beloniformes, Hemiramphidae) from the Lower Sarmatian of the Krasnodar Region |url=https://doi.org/10.1134/S0031030116060034 |journal=Paleontological Journal |language=en |volume=50 |issue=6 |pages=616–622 |doi=10.1134/S0031030116060034 |bibcode=2016PalJ...50..616B |s2cid=89531400 |issn=1555-6174|url-access=subscription }}</ref> Apart from differences in the length of the upper and lower jaws, recent and fossil halfbeaks are distinguished by the fusion of the third pair of upper pharyngeal bones into a plate.<ref name="Lovejoy"/>

==Phylogeny== {{Cladogram|caption=Phylogeny of the halfbeaks. |clades={{clade |1={{clade |1={{clade |label1= |1=Belonidae+Scomberesocidae |label2= |2={{fontcolor|||Zenarchopteridae}} }} |2={{clade |label1= |1={{fontcolor|||'''Hemiramphidae''' }} |label2= |2=Exocoetidae }} }} |label2= |2=Adrianichthyidae }} }} The phylogeny of the halfbeaks is in a state of flux.

On the one hand, there is little question that they are most closely related to three other families of streamlined, surface water fishes: the flyingfishes, needlefishes, and sauries.<ref name="Helfman">{{ cite book|title=The Diversity of Fishes|last=Helfman|first=Gene S.|author2=Collette, Bruce B. |author3=Facey, Douglas E. |publisher=Blackwell Publishing|year=1997|pages=274–276|isbn=978-0-86542-256-8}}</ref> Traditionally, these four families have been taken to comprise the order Beloniformes.<ref name="Helfman"/> The halfbeaks and flyingfishes are considered to form one group, the superfamily Exocoetoidea, and the needlefishes and sauries another, the superfamily Scomberesocoidea.<ref name="Nelson">{{ cite book|title=Fishes of the World|last=Nelson|first=Joseph S.|publisher=John Wiley & Sons, Inc|year=2006|pages=278–280|isbn=978-0-471-25031-9|title-link=Fishes of the World}}</ref>

On the other hand, recent studies have demonstrated that rather than forming a single monophyletic group (a clade), the halfbeak family actually includes a number of lineages ancestral to the flyingfishes and the needlefishes. In other words, as traditionally defined, the halfbeak family is paraphyletic.<ref name="Lovejoy"/>

Within the subfamily Hemiramphinae, the "flying halfbeak" genus ''Oxyporhamphus'' has proved to be particularly problematic; while morphologically closer to the flyingfishes, molecular evidence places it with ''Hemiramphus'' and ''Euleptorhamphus''. Together, these three genera form the sister group to the flyingfish family.<ref name="Lovejoy"/> The other two hemiramphine genera ''Hyporhamphus'' and ''Arrhamphus'' form another clade of less clear placement.<ref name="Lovejoy"/>

Rather than being closely related to the flyingfishes, the subfamily Zenarchopterinae appears to be the sister group of the needlefishes and sauries. This is based on the pharyngeal jaw apparatus, sperm ultrastructure, and molecular evidence. However, this hypothesis has awkward implications for how the morphological evolution of the group is understood, because the fused pharyngeal plate has been considered reliably diagnostic of the halfbeak family.<ref name="Lovejoy"/> Furthermore, the existing theory that because juvenile needlefish pass through a developmental stage where the lower jaw is longer than the upper jaw (the so-called "halfbeak stage") the theory that halfbeaks are paedomorphic needlefish is untenable.<ref name="Lovejoy"/> In fact the unequal lengths of the upper and lower jaws of halfbeaks appears to be the basal condition, with needlefish being relatively derived in comparison.<ref name="Lovejoy">{{cite journal|last=Lovejoy|first=N|author2=Iranpour, M |author3=Collette, B |title=Phylogeny and Jaw Ontogeny of Beloniform Fishes|journal=Integrative and Comparative Biology|volume=44|pages=366–377|year=2004|doi=10.1093/icb/44.5.366|issue=5|pmid=21676722|doi-access=free}}</ref><ref>{{cite journal|last=Boughton|first=D|author2=Collette, B |author3=McCune, A. |title=Heterochrony in Jaw Morphology of Needlefishes (Teleostei: Belonidae)|journal=Systematic Zoology|volume=40|pages=329–352|year=1991|doi=10.2307/2992326|issue=3|jstor=2992326}}</ref>

==Morphology== thumbnail|right|Preserved specimen The halfbeaks are elongated, streamlined fish adapted to living in open water. They can grow to over {{convert|40|cm|in|abbr=on}} in standard length in the case of ''Euleptorhampus viridis''. Their scales are relatively large, cycloid (smooth), and easily detached. No spines are in the fins.<ref name="Collette"/> A distinguishing characteristic is that the third pair of upper pharyngeal bones is anklylosed (fused) into a plate. Halfbeaks are one of several fish families that lack a stomach, all of which possess a pharyngeal jaw apparatus (pharyngeal mill).<ref name="Nelson"/> Most species have an extended lower jaw, at least as juveniles, though this feature may be lost as the fish mature, as with ''Chriodorus'', for example.<ref name="Lovejoy"/>

As is typical for surface-dwelling, open-water fish, most species are silvery, darker above and lighter below, an example of countershading. The tip of the lower jaw is bright red or orange in most species.<ref name="Collette"/>

Halfbeaks carry several adaptations to feeding at the water surface. The eyes and nostrils are at the top of the head, and the upper jaw is mobile, but not the lower jaw. Combined with their streamlined shape and the concentration of fins towards the back (similar to that of a pike), these adaptations allow halfbeaks to locate, catch, and swallow food items very effectively.<ref name="Hoedeman">{{ cite book|last=Hoedeman|first=J.|title=Naturalist's Guide to Freshwater Aquarium Fish|publisher=Elsevier|pages=724–729|year=1974|isbn=978-0-8069-3722-9}}</ref>

==Range and habitat== thumb|250px|right|A marine halfbeak, ''Hemiramphus balao'' Halfbeaks inhabit warm seas, predominantly at the surface, in the Atlantic, Indian, and Pacific Oceans.<ref name="Nelson"/> A few are found in estuaries. Most species of marine halfbeaks are known from continental coastlines, but some extend into the western and central Pacific, and one species (''Hyporhamphus ihi'') is endemic to New Zealand.<ref name="Nelson"/> ''Hemiramphus'' is a worldwide marine genus.<ref name=fishbase/>

==Ecology and behavior==

===Feeding=== Marine halfbeaks are omnivores feeding on algae, marine plants such as seagrasses, plankton, invertebrates such as pteropods and crustaceans, and smaller fishes.<ref>{{ cite journal|last=Randall|first=J|url=http://www.aoml.noaa.gov/general/lib/CREWS/Cleo/PuertoRico/prpdfs/randall-habits.pdf|title=Food habits of reef fishes of the West Indies|journal=Studies in Tropical Oceanography|volume=5|pages=665–847|year=1967|format=PDF Converted to digital format by NOAA, 2004|access-date=2009-06-22}}</ref> For some subtropical species, at least, juveniles are more predatory than adults. Some tropical species feed on animals during the day and plants at night, while other species alternate between carnivory in the summer and herbivory in the winter.<ref name="Tibbetts">{{ cite journal|last=Tibbetts|first=I|author2=Carseldine, L|title=Trophic shifts in three subtropical Australian halfbeaks (Teleostei: Hemiramphidae)|journal=Marine & Freshwater Research|volume=56|pages=925–932|year=2005|doi=10.1071/MF04305|issue=6}}</ref> They are, in turn, eaten by many ecologically and commercially important fish, such as billfish, mackerel, and sharks, so are a key link between trophic levels.<ref name="Mahmoudi">{{ cite web |last=Mahmoudi |first=B |author2=McBride, R |title=A review of Florida's halfbeak bait fishery and halfbeak biology, and a preliminary stock assessment |url=http://research.myfwc.com/images/articles/17405/bally_2002rff_3251.pdf |publisher=Florida Fish and Wildlife Conservation Commission |year=2002 |access-date=2009-06-22 |url-status=dead |archive-url=https://web.archive.org/web/20071025220828/http://research.myfwc.com/images/articles/17405/bally_2002rff_3251.pdf |archive-date=2007-10-25 }}</ref>

===Behavior=== Marine halfbeaks are typically pelagic, schooling, forage fish.<ref name="McGarvey"/><ref name="Stewart"/> The southern sea garfish, ''Hyporhamphus melanochir'' for example, is found in sheltered bays, coastal seas, and estuaries around southern Australia in waters down to {{convert|20|m|ft|abbr=on}}. These fish school near the surface at night, but swim closer to the sea floor during the day, particularly among beds of seagrasses.<ref name="Lyle"/> Genetic analyses of the different subpopulations of the eastern sea garfish ''Hyporhamphus melanochir'' in South Australian coastal waters reveal a small but consistent migration of individuals among them, sufficient to keep them genetically homogeneous.<ref name="McGarvey"/>

Some marine halfbeaks, including ''Euleptorhamphus velox'' and ''E. viridis'', are known for their ability to jump out of the water and glide over the surface for considerable distances, and have consequently sometimes been called flying halfbeaks'.<ref name="fishbase"/><ref>{{FishBase|genus=Euleptorhamphus|species=velox|year=2007|month=Aug}}</ref><ref>{{FishBase|genus=Euleptorhamphus|species=viridis|year=2007|month=Aug}}</ref>

===Reproduction=== Hemiramphidae species are all external fertilizers.<ref name="Nelson"/> They are usually egg-layers and often produce relatively small numbers of fairly large eggs for fish of their size, typically in shallow coastal waters, such as the seagrass meadows of Florida Bay.<ref name="McBride">{{ cite journal|url=http://www.biolbull.org/cgi/content/abstract/204/1/57|title=Reproductive Biology of ''Hemiramphus brasiliensis'' and ''H. balao'' (Hemiramphidae): Maturation, Spawning Frequency, and Fecundity (abstract)|first=Richard S.|last=McBride|author2=Thurman, Paul E.|journal=Biol. Bull.|volume=204|pages=57–67|date=February 2003|doi=10.2307/1543496|pmid=12588745|issue=1|jstor=1543496|s2cid=26249413}}</ref> The eggs of ''Hemiramphus brasiliensis'' and ''H. balao'' are typically {{convert|1.5|-|2.5|mm|in|abbr=on}} in diameter and have attaching filaments. They hatch when they grow to about {{convert|4.8|-|11|mm|in|abbr=on}} in diameter.<ref name="Collette"/> ''Hyporhamphus melanochir'' eggs are slightly larger, around {{convert|2.9|mm|in|abbr=on}} in diameter, and are unusually large when they hatch, being up to {{convert|8.5|mm|in|abbr=on}} in size.<ref name="Lyle"/>

Relatively little is known about the ecology of juvenile marine halfbeaks, though estuarine habitats seem to be favored by at least some species.<ref name="Lyle"/><ref name="McBride"/> The southern sea garfish ''Hyporhamphus melanochir'' grows rapidly at first, attaining a length up to {{convert|30|cm|in|abbr=on}} in the first three years, after which growth slows.<ref name="Lyle"/> This species lives for a maximum age of about 9 years, when the fish reach up to {{convert|40|cm|in|abbr=on}} and weigh about {{convert|0.35|kg|lb|abbr=on}}.<ref name="Lyle"/>

==Relationship to humans==

===Halfbeak fisheries=== Halfbeaks are not a major target for commercial fisheries, though small fisheries for them exist in some places, for example in South Australia, where fisheries target the southern sea garfish (''Hyporhamphus melanochir'').<ref name="McGarvey">{{cite journal |first=R. |last= McGarvey |author2=Fowler, A. J. |author3=Feenstra, J. E. |author4=Jackson, W. B. | title=Garfish (''Hyporhamphus melanochir'') Fishery, Fishery Assessment Report to PIRSA for the Marine Scalefish Fishery Management Committee|journal=SARDI Research Report Series|volume=163 |year=2006 |pages=1–55}}</ref> and the eastern sea garfish (''Hyporhamphus australis'').<ref name="Stewart">{{cite journal |first=John|last= Stewart|title=An observer-based assessment of the estuarine fishery for eastern sea garfish (''Hyporhamphus australis'') in Australia|journal=NSW Department of Primary Industries – Fisheries Final Report Series|volume=87 |year=2007 |pages=1–14}}</ref> Halfbeaks are caught by a variety of methods, including seines and pelagic trawls, dip-netting under lights at night, and haul nets.<ref name="Collette"/> They are used fresh, dried, smoked, or salted,<ref name="Collette"/><ref name="McGarvey"/><ref name="Stewart"/> and they are considered good eating. Even when halfbeaks are targeted by fisheries, though, they tend to be of secondary importance compared with other edible fish species.<ref name="Lyle">{{cite journal |first=J. M.|last= Lyle|author2=Jordan, A. R.|title=Tasmanian Scaleless Fisheries Assessment|journal=Tasmanian Aquaculture and Fisheries Institute Fishery Assessment Reports|volume=1 |year=1988 |pages=1–87}}</ref>

In some localities, significant bait fisheries exist to supply sport fishermen.<ref name="Collette">{{cite web|last=Collette |first=Bruce B. |title=Family Hemiramphidae Gill 1859 – Halfbeaks |url=http://www.calacademy.org/research/ichthyology/annotated/checklists/Hemiramphidae.pdf |publisher=California Academy of Sciences Annotated Checklist of Fishes |issue=22 |date=February 2004 |url-status=dead |archive-url=https://web.archive.org/web/20060623084203/http://www.calacademy.org/research/ichthyology/annotated/checklists/Hemiramphidae.pdf |archive-date=2006-06-23 }}</ref><ref name="Tibbetts"/><ref name="Mahmoudi"/> One study of a bait fishery in Florida that targets ''Hemiramphus brasiliensis'' and ''H. balao'' suggests that despite increases in the size of the fishery, the population is stable and the annual catch is valued at around $500,000.<ref name="Mahmoudi"/><ref>{{cite web|url=http://www.nmfs.noaa.gov/mb/sk/saltonstallken/halfbeak.htm|title=Status and Trends of Florida's Halfbeak Fishery|access-date=2007-02-10}}</ref>

==See also== {{Commons category|Hemiramphidae}} {{Wikispecies|Hemiramphidae}} * USS ''Halfbeak'' - an American submarine named after these fish * USS ''Balao'' - another US submarine (and class) named after a Hemiramphidae species {{Clear}}

==References== {{Reflist|2}}

Category:Beloniformes Category:Hemiramphidae Category:Fish of Hawaii Category:Extant Eocene first appearances Category:Paraphyletic groups