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.

Higher classification of mealybugs (Hemiptera: Coccomorpha) inferred from molecular phylogeny and their endosymbionts

“…In most previous phylogenetic analyses, Putoidae consistently formed a separate clade outside the main cluster of Pseudococcidae (Cook et al, 2002;Downie & Gullan, 2004;Gullan & Cook, 2007;Hodgson, 2002;Hodgson & Hardy, 2013;Vea & Grimaldi, 2016). Here, the type species of Ceroputo Šulc, C. pilosellae Šulc, was placed within Phenacoccinae instead of Putoidae and supports the transfer of Ceroputo to Phenacoccinae from Putoidae (Hardy, Gullan, & Hodgson, 2008;Williams et al 2011).…”

“…The adult females of Rastrococcus have a peculiar formation of the cerarian setae that are likely to support lateral wax filaments (Cox, 1987). The structure and number of cerarii are taxonomically (1978) significant for classifying the mealybugs at various levels (Danzig & Gavrilov-Zimin, 2014;Williams et al 2011). In general, Rastrococcus species have multiple truncated conical setae (ca.…”

Insectos escama (Hemiptera: Coccomorpha) en raíces de café de Norte de Santander y Valle del Cauca, Colombia y descripción de una nueva especie

“…Estos resultados son los esperados, dado que las especies de Pseudococcidae tienen una alta plasticidad biológica que les permite habitar en estructuras aéreas y subterráneas y alimentarse de diferentes tejidos de la planta (McKenzie 1967; Williams y Granara de Willink 1992), de allí que existan cerca de 314 especies asociadas a raíces (McKenzie 1967;Williams 1985;Watson y Cox 1990;Ben-Dov 1994;Gimpel y Miller 1996;Kosztarab 1996;Miller y Arapu 1997;Hara et al 2001;Culik et al 2007;Danzig 2007;Kondo et al 2008b Por otro lado, aunque Coccidae es la tercera familia más diversa de Coccomorpha (Ben-Dov y Hodgson 1997; García-Morales et al 2016), las especies que la componen tienden a alimentarse de estructuras aéreas de la planta (Kosztarab y Kozár 1988), muestra de ello es que solo 32 especies han sido registradas alimentándose de tejido radical (Gill 1988;Kosztarab y Kozár 1988;Ben-Dov 1994;Kosztarab 1996;Kondo y Gullan 2004;Ben-Dov 2012;Kondo 2013;Moghaddam 2013;García-Morales et al 2016). El número bajo de especies de Ortheziidae y Putoidae probablemente se debe a que son familias compuestas por menos especies respecto a Pseudococcidae y Coccidae y todavía existen vacíos en la información de su ecología y biología (Kozár 2004;Williams et al 2011;Vea 2014). Por otra parte, Rhizoecidae tiene como mayor centro de diversidad el Neotrópico, mientras que para Ortheziidae es el Paleartico (Kozár 2004;Kozár y Konczné 2007).…”

Bacterial associates of Orthezia urticae, Matsucoccus pini, and Steingelia gorodetskia - scale insects of archaeoccoid families Ortheziidae, Matsucoccidae, and Steingeliidae (Hemiptera, Coccomorpha)

“…Scale insects are usually divided into two groups: Orthe z ioidea Amyot et Serville, 1843 and Coccoidea Fallen, 1814 which are considered as superfamilies (Koteja 1974; Danzig 1980; Williams and Watson 1990; Morales 1991; Kosztarab 1996; Ben-Dov 2005; Gavrilov-Zimin 2018). Other researchers treat these groups as informal archaeococcoids and neococcoids (Cook et al 2002; Foldi 2005; Hodgson and Foldi 2006; Hardy et al 2008; Kaydan and Kozár 2010; Williams et al 2011; Hodgson 2012, 2014; Hodgson and Hardy 2013). In the present work, we have investigated symbiotic systems of three species of archaeococcoids (= Orthe z ioidea): Orthezia urticae (Linnaeus, 1758), Matsucoccus pini (Green, 1925), and Steingelia gorodetskia Nasonov, 1908.…”