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Mesozoic Fishes 5 – Global Diversity and Evolution, G. Arratia, H.-P. Schultze & M. V. H. Wilson (eds.): pp. 325-362, 21 figs., 2 tabs., 2 apps.
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The first fossil shellear and its implications
for the evolution and divergence of the Kneriidae
(Teleostei: Gonorynchiformes)
Matthew P. DAVIS, Gloria ARRATIA and Thomas M. KAISER
Abstract
A new genus and species of shellear (Gonorynchiformes: Kneriidae), †Mahengichthys singidaensis, is described
from the Eocene Mahenge deposits in Tanzania, Africa. This work represents the first record of a fossil kneriid
gonorynchiform fish. Previously, all gonorynchiform fossils have been attributed to either the families Chanidae
or Gonorynchidae, with some taxa incertae sedis. We explore the phylogenetic position of †Mahengichthys
singidaensis within the gonorynchiforms, utilizing parsimony and maximum likelihood methodologies that
incorporate both morphological and molecular data. Our results indicate that †Mahengichthys singidaensis is a
kneriid gonorynchiform within the tribe Kneriini, which includes the extant genera Kneria and Parakneria. This
phylogenetic work provides a framework for estimating the divergence times of the Kneriidae for the first time
using Bayesian methodology with calibrations that include information regarding extinct kneriids. We infer
that the exclusively freshwater family Kneriidae most likely diverged and diversified during the Cretaceous to
Paleogene in Sub-Saharan Africa, following the continent’s separation from South America.
Introduction
The order Gonorynchiformes includes seven extant and approximately seventeen extinct genera of fishes
that exhibit incredible morphological diversity, with the clade dating back to the Early Cretaceous (FARA
et al. 2010, POYATO-ARIZA et al. 2010a). There are three monophyletic families of gonorynchiform fishes,
including the predominantly marine families Chanidae and Gonorynchidae, and the exclusively freshwater
family Kneriidae distributed throughout Sub-Saharan Africa. While currently there is a general consensus
regarding the monophyly of the order and each family, the relationships of the families to one another
remains controversial, with conflict between molecular (LAVOUÉ et al. 2005, LAVOUÉ et al. 2012) and
morphological (e. g., GRANDE & POYATO-ARIZA 1999, POYATO-ARIZA et al. 2010b) hypotheses. At
present, the majority of previously described extinct genera are attributed to the families Chanidae and
Gonorynchidae, with a few genera recognized as incertae sedis within the order (e. g., †Halecopsis, †Apul-
ichthys). The only family that lacks a fossil representative is the exclusively freshwater family Kneriidae.
In this study we describe the first fossil specimens of an extinct kneriid species collected in the fos-
sil deposits of the ancient maar lake of Mahenge in Tanzania, Africa. The Cenozoic Mahenge deposits
are considered to be one of the most important freshwater fish localities in Africa (GREENWOOD 1974,
MURRAY 2000a, KAISER et al. 2006) due to the excellent preservation of the specimens, and the diver-
sity of fish fauna previously described. Many taxa important to our broader understanding of teleostean
relationships have been described from this locality, including the clupeid †Palaeodenticeps tanganikae
GREENWOOD, 1960, two osteoglossomorphs †Singida jacksonoides GREENWOOD & PATTERSON, 1967,
and †Chauliopareion mahengeense MURRAY & WILSON, 2005, a siluriform from the genus Chrysichthys
(MURRAY 2003a), a characiform †Mahengecharax carrolli MURRAY, 2003b, and various cichlid taxa (e. g.,
GREENWOOD 1960, GREENWOOD & PATTERSON 1967, MURRAY 2000b) including the extinct genus
†Mahengechromis MURRAY, 2000b, which are among the oldest cichlid fossils currently described from
Africa. Additional material from the locality remains to be described, including non-cichlid percomorph
taxa (KAISER et al. 2006).