The Characiformes likely first originated and diversified on the
supercontinent of
West Gondwana (composed of modern Africa and South America) during the
Cretaceous period, though fossils from this time are poorly known. The characiforms had not spread into Africa soon enough to also reach the
land connection between
Africa and Asia. Other fossil teeth date back to the
Cenomanian of
Morocco, but it has been suggested that these teeth may be of early
ginglymodians. Previously, the oldest characiform was assumed to be
Santanichthys of the Early
Cretaceous (
Albian Age) of
Brazil. This presumably marine taxon was used as evidence of characiformes potentially having marine origins. However, more recent studies indicate that
Santanaichthys is likely a basal
otophysan rather than a characiform. Similarly,
Salminops from Spain and
Sorbinicharax from Italy, previously also considered potential marine characiforms, are now thought to have no characiform affinities and are considered indeterminate
teleosts. Given this, there is no paleontological support for characiforms having marine origins. Two other alleged Eocene European characids,
Prohydrocyon Piton, 1938 and
Procharacinus Piton, 1938 from France, lack a
Weberian apparatus and are very likely not characins, and instead may be related to the enigmatic
Thaumaturus.
Eurocharax Gaudant, 1980 is known from a fully-articulated specimen from the
Oligocene of France, and appears to represent an
estuarine taxon, although its
phylogenetic position needs revision.
Fossil taxa • Order
Characiformes • Genus †
Eotexachara Wick, 2021 (
Late Cretaceous of Texas, US) • Genus †
Primuluchara Wick, 2021 (Late Cretaceous of Alberta, Canada and Texas, US)'' • Suborder
Characoidei • Genus †
Tiupampichthys Gayet & Jegu, 2003 (latest Cretaceous to
late Paleocene of Bolivia) • Superfamily
Characoidea • Genus †
Bryconetes Weiss, Malabarba & Malabarba, 2014 (
Late Eocene/
Early Oligocene of Brazil) • Genus †
Paleotetra Weiss, Malabarba & Malabarba, 2012 (Late Eocene/Early Oligocene of Brazil) '', a fossil characiform from the
Oligocene of Brazil Uniquely, Late Cretaceous characiform fossils are found significantly north of their modern distribution. Indeterminate characiform teeth are known from the Santonian of Hungary and
Maastrichtian of France, which have a large, multi-cusped appearance reminiscent of African
alestids. Similarly, two
Campanian freshwater characiform genera,
Primuluchara and
Eotexachara, are known from North America, with
Primuluchara having a very wide distribution across
Laramidia, ranging from Texas to as far north as southern Canada (
Dinosaur Park Formation). It is likely that the warmer conditions of the Late Cretaceous allowed early characins to range farther north than the present day, with African characins colonizing Europe and South American characins colonizing North America. Early characins may have had some level of salt tolerance, allowing for such colonizations to take place. Within their modern distribution, a number of modern South American characin families have their earliest occurrences in the Maastrichtian of Bolivia, with isolated teeth and skeletal elements identifiable to
Acestrorhynchidae,
Characidae, and
Serrasalmidae.
Phylogeny Below is a phylogeny of living Characiformes based on Betancur-Rodriguez
et al. 2017 and Nelson, Grande & Wilson 2016, with the African clades being marked with
Af; }} ==References==