A study of the scale insect genera Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea: Putoidae) and Ceroputo Šulc (Pseudococcidae) with a comparison to Phenacoccus Cockerell (Pseudococcidae)

Abstract

For almost a century, the scale insect genus Puto Signoret (Hemiptera: Sternorrhyncha: Coccoidea) was considered to belong to the family Pseudococcidae (the mealybugs), but recent consensus accords Puto its own family, the Putoidae. This paper reviews the taxonomic history of Puto and family Putoidae, compares the morphology of Puto to that of Ceroputo Šulc and Phenacoccus Cockerell, and reassesses the status of all species that have been placed in Puto to determine whether they belong to the Putoidae or to the Pseudococcidae. For 49 of 57 species that have been placed in Puto, as listed in the online database ScaleNet, we score and tabulate features that are diagnostic for Putoidae and then list all species in their correct family placement. For comparison, we include a few species of Pseudococcidae, namely five species of Phenacoccus, including the type species Phenacoccus aceris (Signoret), and the type species of Ceroputo, C. pilosellae Šulc. We provide revised synonymy lists for Puto and Ceroputo, a brief diagnosis of each genus, synonymy lists and notes for several species for which we suggest recombinations or additional synonyms, or for which we have additional data on morphology. We provide a brief diagnosis of Phenacoccus for comparison with Ceroputo and Puto. As a result of our study, we recognise 47 extant and two fossil species of Puto, and six species of Ceroputo. The New World species Puto mimicus McKenzie and Puto nulliporus McKenzie are transferred to the mealybug genus Ceroputo as Ceroputo mimicus (McKenzie) comb. nov. and Ceroputo nulliporus (McKenzie) comb. nov., respectively, and the Old World species Puto pini Danzig and Puto vaccinii Danzig are recognised as Ceroputo pini (Danzig) comb. rev. and Ceroputo vaccinii (Danzig) comb. rev., respectively, in agreement with Tang (1992). The Old World species Puto graminis Danzig is transferred to Ceroputo as Ceroputo graminis (Danzig) comb. nov. Based only on a study of the literature, the following two names are treated here as junior subjective synonyms of Ceroputo pilosellae: Phenacoccus asteri Takahashi syn. nov. and Puto jarudensis Tang syn. nov. We agree with Tang (1992) that Leococcus erigeroneus Kanda should be treated as a junior subjective synonym of C. pilosellae and thus the genus name Leococcus Kanda, erected for L. erigeroneus and formerly treated as a junior synonym of Puto, is a junior synonym of Ceroputo.