Halfbeak

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Halfbeak
Temporal range: Eocene–Recent
Ballyhoo (
Hemiramphus brasiliensis
), a marine halfbeak
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Actinopterygii
Order: Beloniformes
Superfamily: Exocoetoidea
Family: Hemiramphidae
T. N. Gill
, 1859
Genera

see text

Hemiramphidae is a

viviparous halfbeaks
(family Zenarchopteridae) have often been included in this family.

Though not

predatory fishes, such as billfishes, mackerels, and sharks
.

Taxonomy

In 1758,

type genus.[1] The name comes from the Greek hemi, meaning half, and rhamphos, meaning beak or bill.[2]

There are currently eight genera within the family Hemirampphidae:[3]

Oxyporhamphus micropterus has been considered either a halfbeak or a flyingfish.

This family is primarily

Pacific, and Indian Oceans, though some inhabit estuaries and rivers
.

Evolution

The halfbeaks'

Lower Tertiary.[2] The earliest known halfbeak is “Hemiramphusedwardsi from the Eocene at Monte Bolca, Italy.[4] 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.[5]

Phylogeny

  

Hemiramphidae

  

Exocoetidae

  

Adrianichthyidae

Phylogeny
of the halfbeaks.

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

sauries.[6] Traditionally, these four families have been taken to together comprise the order Beloniformes.[6] The halfbeaks and flyingfishes are considered to form one group, the superfamily Exocoetoidea, and the needlefishes and sauries another, the superfamily Scomberesocoidea.[7]

On the other hand, recent studies have demonstrated that rather than forming a single

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.[5] The other two hemiramphine genera Hyporhamphus and Arrhamphus form another clade of less clear placement.[5]

Rather than being closely related to the flyingfishes, the subfamily Zenarchopterinae appears to be the sister group of the

derived in comparison.[5][8]

Morphology

Preserved specimen

The halfbeaks are elongate, streamlined fish adapted to living in open water. Halfbeaks can grow to over 40 centimeters (16 in) SL in the case of

Chriodorus, for example.[5]

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.[1]

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.[9]

Range and habitat

A marine halfbeak, Hemiramphus balao

Halfbeaks inhabit warm seas, predominantly at the surface, in the

Pacific, and one species (Hyporhamphus ihi) is endemic to New Zealand.[7] Hemiramphus is a worldwide marine genus.[2]

Ecology and behavior

Feeding

Marine halfbeaks are

crustaceans; and smaller fishes.[10] 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.[11] They are in turn eaten by many ecologically and commercially important fish, such as billfish, mackerel, and sharks, and so are a key link between trophic levels.[12]

Behavior

Marine halfbeaks are typically

pelagic schooling forage fish.[13][14] The southern sea garfish Hyporhamphus melanochir for example is found in sheltered bays, coastal seas, estuaries around southern Australia in waters down to a depth of 20 meters (66 ft). These fish school near the surface at night but swim closer to the sea floor during the day, particularly among beds of seagrasses.[15] Genetic analysis of the different sub-populations of the eastern sea garfish Hyporhamphus melanochir in South Australian coastal waters reveals that there is a small but consistent migration of individuals among theme, sufficient to keep them genetically homogeneous.[13]

Some marine halfbeaks, including Euleptorhamphus velox and Euleptorhamphus 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.[2][16][17]

Reproduction

Hemiramphidae species are all external fertilizers.[7] 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.[18] The eggs of Hemiramphus brasiliensis and H. balao are typically 1.5–2.5 mm (0.059–0.098 in) in diameter and have attaching filaments. They hatch when they grow to about 4.8–11 mm (0.19–0.43 in) in diameter.[1] Hyporhamphus melanochir eggs are slightly larger, around 2.9 mm (0.11 in) in diameter, and are unusually large when they hatch, being up to 8.5 mm (0.33 in) in size.[15]

Relatively little is known about the ecology of juvenile marine halfbeaks, though estuarine habitats seem to be favored by at least some species.[15][18] The southern sea garfish Hyporhamphus melanochir grows rapidly at first, attaining a length of up to 30 cm (12 in) in the first three years, after which point growth slows.[15] This species lives for a maximum age of about 9 years, at which point the fish reach up to 40 cm (16 in) and weigh about 0.35 kg (0.77 lb).[15]

Relationship to humans

Halfbeak fisheries

Halfbeaks are not a major target for

trawls, dip-netting under lights at night, and with haul nets.[1] They are utilized fresh, dried, smoked, or salted,[1][13][14] and they are considered good eating. However, even where halfbeaks are targeted by fisheries, they tend to be of secondary importance compared with other edible fish species.[15]

In some localities significant bait fisheries exist to supply sport fishermen.[1][11][12] One study of a bait fishery in Florida that targets Hemiramphus brasiliensis and Hemiramphus 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.[12][19]

See also

References

  1. ^ a b c d e f g Collette, Bruce B. (February 2004). "Family Hemiramphidae Gill 1859 – Halfbeaks" (PDF). California Academy of Sciences Annotated Checklist of Fishes. Archived from the original (PDF) on 2006-06-23.
  2. ^ a b c d Froese, Rainer, and Daniel Pauly, eds. (2007). "Hemiramphidae" in FishBase. Aug 2007 version.
  3. ^ Eschmeyer, William N.; Fricke, Ron & van der Laan, Richard (eds.). "Genera in the family Hemiramphidae". Catalog of Fishes. California Academy of Sciences. Retrieved 18 August 2019.
  4. S2CID 89531400
    .
  5. ^ .
  6. ^ .
  7. ^ .
  8. .
  9. .
  10. NOAA
    , 2004)
    . Studies in Tropical Oceanography. 5: 665–847. Retrieved 2009-06-22.
  11. ^ .
  12. ^ a b c Mahmoudi, B; McBride, R (2002). "A review of Florida's halfbeak bait fishery and halfbeak biology, and a preliminary stock assessment" (PDF). Florida Fish and Wildlife Conservation Commission. Archived from the original (PDF) on 2007-10-25. Retrieved 2009-06-22.
  13. ^ a b c d McGarvey, R.; Fowler, A. J.; Feenstra, J. E.; Jackson, W. B. (2006). "Garfish (Hyporhamphus melanochir) Fishery, Fishery Assessment Report to PIRSA for the Marine Scalefish Fishery Management Committee". SARDI Research Report Series. 163: 1–55.
  14. ^ a b c Stewart, John (2007). "An observer-based assessment of the estuarine fishery for eastern sea garfish (Hyporhamphus australis) in Australia". NSW Department of Primary Industries – Fisheries Final Report Series. 87: 1–14.
  15. ^ a b c d e f Lyle, J. M.; Jordan, A. R. (1988). "Tasmanian Scaleless Fisheries Assessment". Tasmanian Aquaculture and Fisheries Institute Fishery Assessment Reports. 1: 1–87.
  16. ^ Froese, Rainer; Pauly, Daniel (eds.) (2007). "Euleptorhamphus velox" in FishBase. Aug 2007 version.
  17. ^ Froese, Rainer; Pauly, Daniel (eds.) (2007). "Euleptorhamphus viridis" in FishBase. Aug 2007 version.
  18. ^
    S2CID 26249413
    .
  19. ^ "Status and Trends of Florida's Halfbeak Fishery". Retrieved 2007-02-10.