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The genus Geophagus was rediagnosed by Kullander (1986) who restricted it to include only those species with paired caudal extensions to the swimbladder lined by 6-12 epihemal ‘ribs’ plus a greater number of caudal than abdominal vertebrae.
Some former species were moved into the resurrected genus Satanoperca while others, such as the ‘Geophagus‘ brasiliensis and ‘G.‘ steindachneri groups represent distinct groupings still in need of definitive classification.
Geophagus and a number of related genera are often included in the putative subfamily Geophaginae.
Kullander (1998) conducted a morphology-based phylogenetic study in which the neotropical Cichlidae was divided into six subfamilies of which the Geophaginae contained 16 genera divided among three ‘tribes’:
Acarichthyini – Acarichthys and Guianacara. Crenicaratini – Biotoecus, Crenicara, Dicrossus and Mazarunia. Geophagini – Geophagus, Mikrogeophagus, ‘Geophagus‘ brasiliensis group, ‘Geophagus‘ steindachneri group, Gymnogeophagus, Satanoperca, Biotodoma, Apistogramma, Apistogrammoides and Taeniacara.
Later molecular studies by Farias et al. (1999, 2000, 2001) resulted in the additions of Crenicichla and Teleocichla to the Geophaginae, a result supported by López-Fernández et al. (2005) who conducted the most detailed molecular analysis of the grouping to date including 16 of the 18 genera and 30 species.
However their conclusions regarding interrelationships between genera did vary somewhat from previous hypotheses and can be summarised by the following loosely-defined groups:
– a weakly-supported sister group relationship between Acarichthys and Guianacara.
– a well-supported ‘Satanoperca clade‘ comprising Satanoperca, Apistogramma, Apistogrammoides and Taeniacara.
– a ‘big clade‘ with Geophagus, Mikrogeophagus, ‘Geophagus‘ brasiliensis group, ‘Geophagus‘ steindachneri group, Gymnogeophagus, Biotodoma, Crenicara and Dicrossus.
– a ‘crenicarine clade‘ with Biotoecus and Crenicichla.
No representatives of Teleocichla or Mazarunia were included in the study but the former is well-established as sister to Crenicichla while the latter has grouped closely with Dicrossus and Crenicara in earlier works.
The other main conclusions of the paper are confirmation that Geophaginae is a monophyletic group exhibiting strong signs of having undergone rapid adaptive radiation.