R. Zardoya et A. Meyer, COMPLETE MITOCHONDRIAL GENOME SUGGESTS DIAPSID AFFINITIES OF TURTLES, Proceedings of the National Academy of Sciences of the United Statesof America, 95(24), 1998, pp. 14226-14231
Despite more than a century of debate, the evolutionary position of tu
rtles (Testudines) relative to other amniotes (reptiles, birds, and ma
mmals) remains uncertain. One of the major impediments to resolving th
is important evolutionary problem is the highly distinctive and enigma
tic morphology of turtles that led to their traditional placement apar
t from diapsid reptiles as sole descendants of presumably primitive an
apsid reptiles. To address this question, the complete (16,787-bp) mit
ochondrial genome sequence of the African side-necked turtle (Pelomedu
sa subrufa) was determined. This molecule contains several unusual fea
tures: a (TA), microsatellite in the control region, the absence of an
origin of replication for the light strand in the WANCY region of fiv
e tRNA genes, an unusually long noncoding region separating the ND5 an
d ND6 genes, an overlap between ATPase 6 and COIII genes, and the exis
tence of extra nucleotides in ND3 and ND4L putative ORFs. Phylogenetic
analyses of the complete mitochondrial genome sequences supported the
placement of turtles as the sister group of an alligator and chicken
(Archosauria) clade. This result clearly rejects the Haematothermia hy
pothesis (a sister-group relationship between mammals and birds), as w
ell as rejecting the placement of turtles as the most basal living amn
iotes. Moreover, evidence from both complete mitochondrial rRNA genes
supports a sister-group relationship of turtles to Archosauria to the
exclusion of Lepidosauria (tuatara, snakes, and lizards). These result
s challenge the classic view of turtles as the only survivors of prima
ry anapsid reptiles and imply that turtles might have secondarily lost
their skull fenestration.