Cryptovaranoides is not a squamate

Abstract Accurate reconstruction of the timescale of organismal evolution requires knowledge of the placement of extinct representatives among living branches. The fossil record has the capacity to reframe hypotheses of organismal evolution by producing representatives of clades that predate previously known fossils or node ages. Recently, one fossil with the potential to drastically change current consensus surrounding the timescale of reptile diversification was described from Triassic fissure-fill deposits in the United Kingdom. This taxon, Cryptovaranoides microlanius, was originally placed deep within the squamate crown clade, suggesting that many lineages of lizards and snakes must have appeared by the Triassic and implying long ghost lineages that paleontologists and molecular phylogeneticists have failed to detect using all other available data. Our team questioned this identification and instead suggested Cryptovaranoides had unclear affinities to living reptiles, but this alternative interpretation was again challenged by the team that originally described this species. Here, we dissect the morphological character codings used to support a crown squamate affinity for Cryptovaranoides microlanius and illustrate empirical problems with analyses that hypothesize this taxon to be a crown squamate. Our analyses emphasize the importance of stringency in constructing hypodigms of fossils, particularly when they may be key for proper time calibration of the Tree of Life.