Valid Names Results

Puto Signoret, 1876 (Putoidae)

Nomenclatural History

• Putonia Signoret 1875c: 341. Type species: Putonia antennata Signoret by monotypy . junior homonym Notes: Homonym of Putonia Stal, 1872, in Heteroptera..
• Puto Signoret 1876a: 394. . replacement name
• Macrocerococcus Leonardi 1907b: 151. Type species: Macrocerococcus superbus Leonardi by original designation . junior synonym (discovered by Danzig1980b: 110) Notes: A subjective synonym of Puto.
• Ceroputo Ferris 1918d: 60;. . incorrect synonymy (discovered by Tang1992
• Leococcus Kawai 1980: 96;. . incorrect synonymy

Remarks

• Systematics: The name Puto, proposed by Signoret (1876), was a replacement name for the occupied name Putonia Signoret, a genus described for the single species Putonia antennata Signoret from France by Signoret (1875). Females and immature males of Puto resemble mealybugs, family Pseudococcidae, and especially some species of Phenacoccus Cockerell. Adult females of Phenacoccus and Puto are similar in possessing 18 basic pairs of cerarii, usually 9-segmented antennae, ostioles, and often a circulus. For almost 100 years, species of Puto were included among the Pseudococcidae. However, it has long been known that adult males of Puto possess a row of unicorneal eyes encircling the head plus a pair of ocelli, whereas adult males of species of Pseudococcidae have only a dorsal and ventral pair of unicorneal eyes and a pair of ocelli. Beardsley (1969) was the first to place Puto in its own family, Putoidae, based on male features, and this action has gained wide recognition (e.g., Hodgson & Foldi, 2006; Gullan & Cook, 2007; Williams, et al., 2011). Because this new family name did not describe the taxon sufficiently, it is considered a nomen nudum and the correct author of the family is Tang 1992. (Gavrilov-Zimin & Danzig 2012). Gavrilov-Zimin and Danzig (2012) provide evidence that Puto should not be included in a separate family but should be part of the Pseudococcidae and should include 2 subgenera Puto and Ceroputo. In 2014, they created several morphological groups of genera (g/g) within the Pseudococcidae. One of these groups is g/g Puto. "In addition to the genus Puto itself with its two subgenera (they) include also in the group monotypic genera Trimerococcus (North Africa) and Eastia (South Africa) and (the) large tropical genus Rastrococcus Ferris, 1954, some species of which are known also from the utmost South-East of (the) Palaearctic (South of China)" They include a section on pages 45-59 concerning their opinion of the taxonomic position of the genus Puto.

  Phylogenetic analyses based on nucleotide sequence data have shown that the genus Puto does not belong to the Pseudococcidae (Cook et al. 2002; Gullan & Cook 2007; Hardy et al. 2008; Choi & Lee 2022) but is a separate archaeococcoid family, Putoidae, as indicated by Williams et al. (2001). Its family status is strongly supported also by morphological features of the adult males (Hodgson & Foldi 2006; Hodgson & Hardy 2013; Hodgson 2020). Choi & Lee (2022) recovered the clade of Puto species placed outside Xenococcidae + Rhizoecidae + Rastrococcinae + Phenacoccinae + Pseudococcinae. Their study supported Putoidae being a separate family from the genus Ceroputo and other Pseudococcidae based on the molecular and morphological evidence plus the possession of a particular/different type of endosymbionts.

• Structure: Adult female: eye height usually as great as length of first antennal segment; antennae usually with 9 segments, rarely with 8; antennal intersegmental sensilla present between segments III-IV, IV-V and VI-VII; each surface of trochanter with 2-5 (usually 3 or 4) campaniform sensilla; claw usually with pair of basal spurs; claw digitules often capitate but tarsal digitules almost never capitate; at least 18 pairs of cerarii on sclerotised plates, if more pairs present, increase due to division of cerarii on certain segments; long tubular ducts almost always present on frons (anterior to mouthparts); multilocular pores present on venter (except in P. peyerinhoffi); quinquelocular pores absent (except in P. peyerinhoffi); trilocular pores usually of 3 sizes, ventral pores smallest, dorsal pores noticeably larger, cerarian pores slightly larger than dorsal pores. Third-instar female: resembles adult female but with 8-segmented antennae, 2-3 campaniform sensilla on each surface of trochanter, fewer pores and ducts and a non-functional vulva. First-instar nymph: 7-segmented antennae; multilocular pores with >5 loculi. Adult male: row of 8-15 (usually 14) eyes surrounding head, plus a pair of lateral ocelli; with one pair of lateral filaments near apex of abdomen (each filament from glandular pouch on each side of abdominal segment VIII); penial sheath 1-segmented, apex of aedeagus either bifid or simple. (Williams, et al., 2011).
• General Remarks: Definition and characters by Borchsenius (1949), Ferris (1950b), McKenzie (1967), Miller & McKenzie (1973), Tereznikova (1975), Danzig (1980b), Kosztarab & Kozár (1988F), Williams & Granara de Willink (1992), Marotta & Tranfaglia (1993), Kosztarab (1996) and by Williams (2004a). Redescrption and analysis in Williams et al. (2011).

Keys

• PowellMi2024: pp.302-303 ( Life Stages ) [Puto species instars]
• Hodgso2020: pp.19-22 ( Adult (M) ) [Neococcoid higher taxa]
• DanzigGa2014: pp.138 ( Adult (F) ) [Key to genera of g/g Puto]
• Willia2004a: pp.39 ( Adult (F) ) [Genera of southern Asia]
• MarottTr1993: pp.174 ( Adult (F) ) [Central Europe and Mediterranean Basin]
• Tang1992: pp.384 ( Adult (F) ) [Palaearctic & Oriental regions]
• WilliaGr1992: pp.41 ( Adult (F) ) [Central and South America]
• WilliaGr1992: pp.468-469 ( Adult (F) ) [Central and South America]
• Danzig1988: pp.697 ( Adult (F) ) [ Far East of the Russia]
• Danzig1988: pp.698-699 ( Adult (F) ) [Species Far East of the USSR]
• KosztaKo1988F: pp.136 ( Adult (F) ) [Central Europe ]
• Matile1985: pp.140 ( Adult (F) ) [Species South America]
• Danzig1980b: pp.112-113 ( Adult (F) ) [Far East Russia]
• Danzig1978b: pp.124-125 ( Adult (F) ) [Far East of the USSR]
• Terezn1975: pp.160 ( Adult (F) ) [Ukraine & Palaearctic region]
• Terezn1975: pp.240 ( Adult (F) ) [Ukraine]
• TerGri1973: pp.38 ( Adult (F) ) [Armenia]
• AfifiKo1967: pp.10 ( Adult (F) ) [World]
• McKenz1967: pp.327-329 ( Adult (F) ) [North America]
• McKenz1967: pp.45 ( Adult (F) ) [North America]
• McKenz1961: pp.32 ( Adult (F) ) [North America]
• McKenz1960: pp.690 ( Adult (F) ) [North America ]
• McKenz1960: pp.730-731 ( Adult (F) ) [North America]
• Ferris1953a: pp.282 ( Adult (F) ) [North America]
• Ferris1950b: pp.21 ( Adult (F) ) [North America]
• Borchs1949: pp. ( Adult (F) ) [USSR]
• Borchs1949: pp.286 ( Adult (F) ) [Palaearctic region]
• Borchs1949: pp.89 ( Adult (F) ) [Palaearctic region]
• Borchs1948b: pp.33 ( Adult (F) ) [six species of USSR]

Associated References

• Afifi1968: taxonomy, pp. 155
• BenDov1994: catalog, pp. 75, 213, 220, 417-41
• Borchs1949: description, taxonomy, pp. 285-286, 289-291
• Danzig1980b: description, taxonomy, pp. 109-113
• DanzigGa2014: description, distribution, host, illustration, structure, taxonomy, pp. 137-178
• Fernal1903b: catalog, pp. 89, 94
• Ferris1950b: description, taxonomy, pp. 190-192
• Ferris1953a: taxonomy, pp. 423
• FoldiKo2006: taxonomy, pp. 309
• Gavril2021b: reproduction, pp. 245
• GavrilDa2012: phylogeny, taxonomy, pp. 97-111
• GullanCo2001: phylogeny, taxonomy, pp. 91-99
• HodgsoFo2005: phylogeny, taxonomy, pp. 35-48
• HodgsoFo2006: taxonomy, pp. 218
• Koszta1996: description, taxonomy, pp. 77, 188-189
• KosztaKo1988F: description, taxonomy, pp. 135-136
• Koteja2000c: taxonomy, pp. 207
• Koteja2001: taxonomy, pp. 48
• KwonDaPa2003: taxonomy, pp. 118
• LinKoGu2013: molecular data, phylogeny, pp. 257
• MarottTr1985: description, taxonomy, pp. 216-218
• MarottTr1993: description, taxonomy, pp. 171-173
• McKenz1967: description, taxonomy, pp. 326-329
• MillerMc1973: description, taxonomy, pp. 515-516
• MoghadWa2022: taxonomy, pp. 140 , 142
• MorrisMo1966: taxonomy, pp. 33, 106, 113, 170
• Signor1875c: taxonomy, pp. 341
• Signor1876a: taxonomy, pp. 394
• Tao1999: taxonomy, pp. 10-11, 26
• Terezn1975: description, taxonomy, pp. 238-240
• Willia2004a: description, taxonomy, pp. 711
• WilliaGr1992: description, taxonomy, pp. 465-469
• WilliaGuMi2011: description, distribution, illustration, taxonomy, pp. 1-22

51 Species