“…Even though these studies have used data from a range of methods, including behavior, morphometrics, karyotype, isozymes, pheromone, cuticular hydrocarbons, mtDNA and reproductive studies, they lack a comprehensive sampling across the species’ range and fail to fully integrate molecular and phenotypic data, which complicates their use in species identification. An indication of the complex taxonomic questions in this genus is that single-gene molecular phylogenies based on mtDNA ( McPheron et al, 1999 ; Smith-Caldas et al, 2001 ) or nuclear genes ( Barr et al, 2005 ; Ruiz et al, 2007 ; Sarno et al, 2010 ; Gonçalves et al, 2013 ), and even using multiple genes ( Mengual et al, 2017 ) have failed to identify species-specific fixed differences among the most closely related taxa, leading some to propose a multidisciplinary approach ( Dias et al, 2016 ).…”