Evolution and classification

Paleontology

The order Gasterosteiformes appears to represent an early but highly specialized branch of the Acanthopterygii. Its evolution has been traced through limited paleontological data. Fossil sticklebacks occur in rock strata from the Miocene Epoch (23–5.3 million years ago) and are moderately abundant in strata formed throughout the Paleogene Period (65.5–23 million years ago). Most of the order occurs in fossil remains dated to the Eocene and Oligocene epochs (55.8–23 million years ago) in the area of Monte Bolca near Verona, Italy. Deposits there and in Sumatra include representatives from the family Protosyngnathidae, an extinct group related to the tubesnouts. Members of a closely related family of extinct scaleless fishes (Urosphenidae), trumpetfish species, and a small species of cornetfish have also been found there. In addition, shrimpfishes are found within these strata as well as in Oligocene deposits in various parts of Europe. Snipefishes are represented by an extinct genus, as are ghost pipefishes. Pipefishes occur in Miocene deposits in Sicily. Syngnathus and Calamostoma, pipefish genera that have a close relationship to true seahorses, have been found in rocks from both the Paleogene and Neogene periods. Fossil seahorses and pegasids are unknown.

Distinguishing taxonomic features

The Gasterosteiformes are classified mainly on the basis of general body form, the structure and distribution of scales or body plates, fin form and position, and the structure of the skeleton and its individual parts.

Annotated classification

The classification here is a combination of J.S. Nelson (Fishes of the World, 4th ed., 2006) and the online taxonomic database FishBase (2021).

• Order Gasterosteiformes

  Frequently with strong spines in dorsal and pelvic fins, spines absent in some; snout often elongated; body often with dermal plates. Length from about 2 to 200 cm (0.8 to 79 inches). 11 families, 374 species. Marine, brackish, and freshwater; widely distributed. Eocene to present.

  • • Family Gasterosteidae (sticklebacks)

      Jaws short, armed with sharp teeth; body quite fusiform (tapered at both ends); body plates may be absent or may vary in number; body length to about 18 cm (7 inches). 5 genera (Gasterosteus, Spinachia, Apeltes, Pungitius, and Culaea), 20 species. Fresh, brackish, and marine waters of Northern Hemisphere.

    • Family Aulorhynchidae (tubesnouts)

      Body elongated, slender, and cylindrical; snout long, upper jaw hinged. 2 genera (Aulichthys and Aulorhynchus), 2 species; northern Pacific Ocean.

    • Family Indostomidae (indostomid or paradox fish)

      Body elongated, covered with bony rings; teeth minute, gills lobe-shaped, eyes large. 3 species; South Asian and Southeast Asian streams.

  • • Family Aulostomidae (trumpetfishes)

      Body elongated and compressed sideways; jaws short and weak, teeth minute; dorsal spines numerous; length to about 80 cm (31 inches). 1 genus (Aulostomus), about 3 species; tropical seas.

    • Family Fistulariidae (cornetfishes)

      Similar in appearance to Aulostomidae; no scales, bony plates imbedded in skin; dorsal spines absent; backbone extends through caudal fin as a central filament. Maximum length 200 cm (79 inches). 1 genus (Fistularia), about 5 species; tropical seas.

    • Family Centriscidae (shrimpfishes)

      Body small, jaws toothless, scales absent, back covered by transparent plates. 2 genera (Aeoliscus and Centriscus), 12 species; shallow waters of Indian and Pacific oceans.

    • Family Macrorhamphosidae (snipefishes or bellows fish)

      Snout tubular, jaws short, body rather short and deep; in profile, shaped like a bellows. 3 genera (Centriscops, Macroramphosus, and Notopogon), 8 species; temperate and tropical seas.

    • Family Solenostomidae (ghost pipefishes)

      Snout tubular, mouth small; body short, with spiny dorsal and ventral fins; bony plates united to form body rings. 1 genus (Solenostomus), 5 to 6 species; tropical Indo-Pacific waters.

    • Family Syngnathidae (pipefishes, seahorses, sea dragons, and pipehorses)

      Pipefishes long and slender, snout tipped with small mouth; dorsal and pectoral fins spineless, ventral fins absent; 59 genera, about 307 species; widely distributed in shallow tropical seas. Seahorses with head bent downward in horselike relation to body; tail prehensile; bony rings instead of scales; 1 genus (Hippocampus), about 50 species; widely distributed, marine. Sea dragons have deep laterally flattened bodies with elaborate, permanent, leaflike appendages; 2 genera (Phyllopteryx and Phycodurus), 2 species.

    • Family Pegasidae (pegasids or dragonfish)

      Toothless mouth is not terminal but lies under the head, overhung by a rostrum (snout), often adorned with spines. Small; body protected by knobby armoured plates; posterior portion rather elongated, square to rectangular in cross section; bearing small dorsal fin; spines along either side of this region may be absent or developed to varying degrees. Pectoral fins form expansive fans on either side; ventral fins reduced to fingerlike rays for crawling along the seabed. Maximum length 18 cm (7 inches). 2 genera (Pegasus and Eurypegasus), 5 to 6 species. Marine; temperate coastal Indo-West Pacific; salt and brackish waters at depths of up to 150 metres (about 500 feet).

    • Family Hypoptychidae (sand lances)

      Body elongate; scutes, scales, and spines absent. Dorsal and anal fins placed posteriorly; pectoral fins and caudal fins; all have soft rays. Maximum length about 8.5 cm (3.5 inches). 2 genera, 2 species (Hypoptychus dybowskii and Aulichthys japonicus). Marine; found from around Japan and Korea to the Sea of Okhotsk.

Critical appraisal

Gasterosteiform classification has not been solidified. The pegasids have been tentatively placed in their own family (Pegasidae) in this classification; however, some authorities group the pegasids with either the sticklebacks (family Gasterosteidae) or the pipefishes and their allies (family Syngnathidae). In some classifications the pegasids have been given their own order (Pegasiformes). Furthermore, some authorities place the families Aulostomidae, Centriscidae, Fistulariidae, Solenostomidae, and Syngnathidae in the order Syngnathiformes.