higher-level (tribal and above) phylogenetic analyses of the family based on character sets are a relatively recent phenomenon,starting with Brower’s (2000) phylogenetic analysis of 103 nymphalid genera based on 378 base pairs from the nuclear gene wingless. de Jong et al. (1996) included representatives from all recognized subfamilies but, apart from consistently placing Liby-theinae as the sister of the remaining Nymphalidae, were unable to resolve interrelationships between them. Ackery et al. (1999) noted in that respect that (p. 287) “most relationships within and between the various groups remain obscure”.