sentence: Jaltomata species also vary in mature fruit color , including species that have either purple , red , orange , or green fruit at maturity ; this fruit color variation appears to characterize three major subclades within the genus as separate dark purple - , red - , and orange-fruited clades ( Miller et al. 2011 ; S ä rkinen et al. 2013 ; Wu et al. 2018 ) .