Key Words: Reptilia; Amphisbaenia; Bipes; phylogeny; mitochondrial DNA; genomics; limb evolution; biogeography; Pangaea.
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INTRODUCTIONAmphisbaenians are fossorial reptiles that inhabit tropical and semitropical regions of the world (Fig. 1). The vast majority of species occur on land masses of Gondwanan origin in South America, Africa and Arabia. However, two of the four families are restricted to regions associated with Laurasia, the Bipedidae, with three species in Mexico (Fig. 2), and the Rhineuridae, with a single species in Florida. These taxa have pleurodont dentition (Gans, 1968) and were originally placed in the family Amphisbaenidae which now primarily includes taxa distributed in Gondwanan regions of South America and Africa (Gans, 1978). The fourth family of Amphisbaenia, the Trogonophidae, is the only one to have acrodont dentition and is restricted to Gondwanan plates of North Africa, Arabia and the island of Socotra. By superimposing a phylogeny of these groups on their distribution relative to the breakup of these major land masses, it is possible to test hypotheses about their origins. For example, monophyly of taxa found in Laurasia or Gondwana may reflect their isolation stemming from the breakup of Pangaea 200 million years before present (MYBP), (Fig. 3C, D).A phylogeny of Amphisbaenia is also important for reconstructing patterns of morphological change. Among the four families of Amphisbaenia, all taxa are limbless except the Bipedidae, which contains reptiles with two forelimbs. If gradual reduction in limbs were to have occurred from an ancestor with both hind and forelimbs, then the Bipedidae, with its presumed intermediate state, would be thought to be the sister taxon to the remaining families as has been suggested by morphological data (Kearney, 2003; Fig. 3B).Within the Bipedidae, species are variable for the number of digits on a limb (Papenfuss, 1982). The Baja California taxon, Bipes biporus, always has five digits on each limb. In mainland Mexico, the more northern taxon, B. canaliculatus, is variable, with either four or five 4 digits, while the more southern species, B. tridactylus, always has three digits on each limb. If digits are being gradually lost from an ancestor with five digits then B. biporus is predicted to be the sister taxon to the other two species (Fig. 3E). This phylogenetic hypothesis is consistent with an initial splitting of Bipes by the separation of peninsular Baja California from the west coast of Mexico (12-14 MYBP;Ferrari, 1995), with the two mainland species later diverging.Alternatively, tectonic activity along the west coast of Mexico in the Cretaceous to Paleocene (Ferrari, 1995) may have isolated the more northern taxa, B. biporus and B. canaliculatus, from B. tridactylus to the south (Fig. 3F).Mitochondrial gene order is a highly robust phylogenetic character for reconstructing evolutionary relationships among animals (Boore, 1999;Boore and Brown 1998;Macey et al., 1997aMacey et al., , 2000. Although no character can be asserted to be completely ...