Phylogenetic systematics and historical biogeography of the Neotropical electric fish Gymnotus (Teleostei: Gymnotidae)

Abstract

Phylogenetic interrelationships of the Neotropical electric fish genus Gymnotus are documented from comparative study of phenotypic data. A data matrix was compiled of 113 phenotypic characters for 40 taxa, including 31 recognized Gymnotus species, six allopatric populations of G. carapo, two allopatric populations of G. coropinae, and three gymno‐tiform outgroups. MP analysis yielded 15 trees of equal length, the strict consensus of which is presented as a working hypothesis of Gymnotus interrelationships. Diagnoses are presented for 26 clades, including three species groups; the G. cylindricus group with two species restricted to Middle America, the G. pantherinus group with 12 species in South America, and the G. carapo group with 16 species in South America. The basal division of Gymnotus is between clades endemic to Middle and South America. Both the G. pantherinus and G. carapo groups include trans‐Andean sister‐taxon pairs, suggesting a minimum date for the origins of these groups in the late Middle Miocene (c. 12 Ma.). The geographically widespread species G. carapo is paraphyletic. Analysis of character state evolution shows characters of external morphology are more phylogenetically plastic and provide more phylogenetic information in recent branches than do characters of internal morphology, which themselves provide the more information in deeper branches. Nine regional species assemblages of Gymnotus are recognized, none of which is monophyletic. There are at least two independent origins of Gymnotus species in sediment rich, high conductivity, perennially hypoxic whitewater floodplains (varzea´) derived from an ancestral condition of being restricted to low conductivity non‐floodplain (terra firme) black and clearwater rivers and streams. These phylogenetic, biogeographic and ecological patterns suggest a lengthy and complex history involving numerous instances of speciation, extinction, migration and coexistence in sympatry. Evolution in Gymnotus has been a continent‐wide phenomenon; i.e. Amazonian species richness is not a consequence of strictly Amazonian processes. These patterns are similar to those of other highly diverse groups of Neotropical fishes and do not resemble those of monophyletic, rapidly generated species flocks.

Keywords:

• biodiversity
• character evolution
• cladistics
• classification
• comparative morphology
• diversification
• historical ecology
• Gymnotiformes
• paraphyletic species
• várzea

Notes

Corresponding author. Now at Department of Biology, University of Louisiana, Lafayette, LA 70504–42451, USA. Email: [email protected]