News                        More
Recent Publications        More
Nomenclature                        More

Introduction to Cypriniformes

Eschmeyer’s (1998 et seq.) Catalog of Fishes includes 32,131 valid species of extant fishes (as of 01/12/12), which is about 50% of the approximately 62,275 species of vertebrates (Nelson 2006, Pennisi 2009). About 41%, or 13,180, of fish species live in fresh water (Berra 2007), which is extraordinary given that fresh water constitutes less than 0.01% of the volume of water on Earth (97% is oceanic; Horn 1972). Furthermore, thousands of species of freshwater fishes remain to be described, most of them in the tropics (Lundberg, et al. 2000).
About 72% of all freshwater fish species (9,254) belong to a single teleost clade, the Otophysi. Otophysi includes four orders, Siluriformes (catfishes) with 3,500 species, Cypriniformes (minnows, loaches and suckers) with 4,000 species, Characiformes (characins) with 1,900 species, and Gymnotiformes (knifefishes) with 160 species.

With 3,862 species, Cypriniformes is the most diverse monophyletic order of vertebrates. Perciformes (perches, basses, cichlids, gobies, etc.) is larger with 9,293 species, but is paraphyletic (Johnson & Patterson 1993; Springer & Johnson 2004). About 1 in 3 (30%) species of freshwater fishes, 1 in 8 fishes (12%), and 1 in 17 vertebrates (6%), is a cypriniform. Cypriniforms are most diverse in the least-explored parts of the tropics, and we estimate the number of species to be much larger than now recognized, closer to 4,500-5,000.

Monophyly of Otophysi, dating from the Eocene (Patterson 1984), is well established by the unique presence of the Weberian ossicles, a modification of the first 4 or 5 vertebrae into moveable bones connecting the inner ear to the gas bladder (Rosen & Greenwood 1970, Fink & Fink 1981, 1996, Chardon & Vandewalle 1991, Fink, et al. 1984). Cypriniformes is diagnosed by the absence of teeth on the jaws and palate, a kinethmoid – unique bone allowing for protrusion of jaws, absence of an adipose fin (present in some cobitoids), three branchiostegal rays, usually scaleless head, and several other skeletal synapomorphies (Fink & Fink 1981, Conway et al. in press, Chen et al. 2009, Mayden & Chen in press). Classification within the order, however, is in a state of flux given that the traditional classification essentially lacked any demonstrable justification, and CToL is providing the framework on which a new classification will be based ((Fig. 2); papers cited below). A new CToL classification will be infused into this PBI when completed.


Eschmeyer, W.N. and Fricke, R. (eds.) 1998 et seq. Catalog of Fishes electronic version (9 September 2009). FAO. 2006. Yearbook of Fishery and Aquaculture Statistics. Food and Agriculture Organization of the United Nations.

Nelson, J. S. 2006. Fishes of the World, 4th edition. John Wiley & Sons, New York.

Pennisi, E. 2009. No genome left behind. Science 326:794-795.

Berra, T. M. 2007. Freshwater Fish Distribution. University of Chicago Press.

Lundberg, J.G., M. Kottelat, G.R. Smith, M. Stiassny, and T. Gill. 2000. So many fishes,so little time: An overview of recent ichthyological discoveries in fresh waters. Annals of the Missouri Botanical Gardens 87: 26-62.

Johnson, G. D. and C. Patterson 1993. Percomorph phylogeny: a survey of acanthomorphs and a new proposal. Bull. Mar. Sci. 52:554-626.

Springer, V. G. and G. D. Johnson. 2004. Study of the dorsal gill-arch musculature of teleostome fishes, with special reference to the Actinopterygii. Bulletin of the Biological Society of Washington 11:1-260.

Patterson, C. 1984. Chanoides, a marine Eocene otophysan fish (Teleostei: Ostariophysi). Journal of Vertebrate Paleontology 4:430-456.

Rosen, D. E. and P. H. Greenwood. 1970. Origin of the Weberian apparatus and the relationships of the ostariophysan and gonorynchiform fishes. American Museum Novitates 2428.

Fink, S. V. and W. L. Fink. 1981. Interrelationships of the ostariophysan fishes (Teleostei). Zoological Journal of the Linnean Society 72:297-353.

Fink, S. V. and W. L. Fink. 1996. Interrelationships of ostariophysan fishes (Teleostei), pp. 209-249 in: M. L. J. Stiassny, L. R. Parenti, and G. D. Johnson (eds.) Interrelationships of Fishes. Academic Press, San Diego.

Chardon, M. and P. Vandewalle. 1991. Acoutico-lateralis system. Pp. 332-352 in I. J. Winfield and J. S. Nelson (Eds.), Cyprinid fishes: systematics, biology and exploitation. Chapman and Hall, London.

Fink, S. V., P. H. Greenwood, and W. L. Fink. 1984. A critique of recent work on fossil ostariophysan fishes. Copeia 1984:1033-1041.

Conway, K. W., M.V. Hirt, L. Yang, R. L. Mayden & A. M. Simons. In press. Cypriniformes: Systematics and Paleontology. In: H-P, Schultz, M. V. H. Wilson & J. S. Nelson (eds):Origin and Phylogenetic Interrelationships of Teleosts. Verlag Dr. Freidrich Feil, München.

Chen, W.-J., L. Lheknim, and R. L. Mayden. 2009. Molecular systematics of the Cobitoidea(Teleostei: Cypriniformes) revisited and phylogenetic position of enigmatic loach Ellopostoma Vaillant 1902: Evidence from six nuclear genes. Journal of Fish Biology 75:9 (December 2009).

Mayden, R. L., and W.-J. Chen. In press. The world’s smallest vertebrate species of the Genus Paedocypris: A new family of freshwater fishes and the sister group to the world’s most diverse clade of freshwater fishes (Teleostei: Cypriniformes). Molecular Phylogenetics and Evolution.


Please install Flash® and turn on Javascript.

If you encounter any security errors loading Monoslideshow, please update your security settings by adding the Monoslideshow folder to the trusted locations.

© 2010 ACSI & CToL

This is the primary page opening to two very large research initiatives on Cypriniformes fishes: the All Cypriniformes Species Inventory (ACSI-2), funded by the U.S. National Science Foundation DEB-1022720 (Dr. Lawrence M. Page, University of Florida), DEB-1023403 (Dr. Jonathan Armbruster, Auburn University), and DEB-1021840 (Dr. Richard L. Mayden, Saint Louis University )and The Cypriniformes Tree of Life (CToL), funded by the U.S. National Science Foundation EF-0431326 (Drs. R. L. Mayden, R. M. Wood, and N. Aspinwall). You can learn more about both of these international-scale studies using the links available on this site.