Foldi, Imre, 2009, Archaeococcoid scale insects (Hemiptera: Coccoidea) from the tropical high mountains of the Andean Cordillera, South America, Zootaxa 2300, pp. 1-38: 4-6

publication ID 10.5281/zenodo.191685

persistent identifier

treatment provided by


scientific name



The family Monophlebidae   in the Neotropical region

The Monophlebidae   is a large family with a worldwide distribution. The classification of the Monophlebinae (now Monophlebidae   ) was first established by Morrison (1927), and discussed further in his remarkable monograph ( Morrison, 1928). Since then there have been additional significant studies, the latest by Unruh & Gullan (2008 a, 2008 b), and Williams & Gullan (2008).

Adult female monophlebids are usually rather large, and important characteristic at all growth stages is their mobility, due to the presence of well-developed brown or black legs. The adult females are characterised by the following morphological features: antennae 7–11 segmented; ventral cicatrices present on the abdomen, usually with 1 or 3 posterior to the vulva, but in some cases many more; multilocular pores often numerous, but rather variable in the structure and configuration of the loculi; multilocular pores also sometimes in groups, forming either an ovisac pore band or marsupial pore band; hairs, hair-like setae, flagellate setae, spiniform setae and spines present or absent; thoracic spiracles usually without pores in atrium; abdominal spiracles varying in number from 2 to 7 pairs, each with or without pores in atrium; anal opening dorsal, with a simple anal tube either with only a sclerotised ring or also with pores at inner end. Identification of monophlebid species is often very difficult. Therefore, for taxonomic purposes, the simultaneous description of the adult female and male, and the first-instar nymph of a given species helps to establish the best characters for its identification and those of related species (Foldi, 2006; Foldi & Watson, 2001).

The taxonomy of the Monophlebidae   in the Neotropical region has been greatly modified by the recent major revision of the tribe Iceryini ( Unruh & Gullan, 2008 a), which is the largest and the most widely distributed tribe in the family (Foldi, 2001; Ben-Dov, 2005). Unruh & Gullan (2008 a) reconstructed the phylogeny of the Iceryini on the basis of molecular data and established numerous new combinations, new synonymies, considered several species to be incertae sedis, and designated lectotypes. They recognised 74 species in the tribe Iceryini belonging to only five genera: Crypticerya Cockerell   , Echinicerya Morrison   , Gigantococcus Pesson & Bielenin   , Gueriniella Fernald   and Icerya Signoret. Unruh & Gullan   (2008 a) redescribed and illustrated the type species of each genus, and provided a key to the genera based on adult females. In addition, Unruh & Gullan (2008 b) published an identification guide to all species in the tribe Iceryini, and the genus Crypticerya   has been revised recently by Unruh (2008). Recently, Kondo & Unruh (2009) described a polyphagus new species, Crypticerya multicicatrices Kondo & Unruh   , from Colombia, with a key to the adult females of South American iceryine species.

The fauna of the Neotropical region now includes 33 known monophlebid species in 7 genera and 2 tribes: 1. Tribe Iceryini: Crypticerya Cockerell   , Echinicerya Morrison   and Icerya Signoret   ; and 2. Tribe Llaveiini: Laurencella Foldi   , Llaveia (Llave)   , Neohodgsonius (Foldi)   and Protortonia Townsend   (Foldi, 2001, Ben-Dov, 2005, Williams & Gullan, 2008, Unruh & Gullan, 2008 a, b). The record of the tribe Monophlebini in the Neotropics is based on Monophlebus niveus Hempel   from Brazil ( Hempel, 1920) and, although the identity of this species has not been established, it is unlikely to be congeneric with the type species of Monophlebus Guérin-Méneville   , M. atripennis Burmeister   from Java. The largest of these genera is Crypticerya   , which now includes 16 species in the Neotropical region excluding the Nearctic part of Mexico: 1. C. abrahami (Newstead)   from Venezuela and Guyana; 2. C. brasiliensis (Hempel)   from Argentina, Brazil, Colombia, Panama and Peru; 3. C. flava (Hempel)   from Brazil; 4. C. flocculosa (Hempel)   from Brazil; 5. C. genistae (Hempel)   from Brazil and recently spread to the Caribben and southeastern USA ( Hodges et al., 2008); 6. C. luederwaldti (Hempel)   from Brazil; 7. C. minima (Morrison)   from Argentina; 8. C. montserratensis (Riley & Howard)   from Colombia, Grenada, Mexico, Montserrat, Panama and Puerto Rico; 9. C. multicicatrices Kondo & Unruh   from Colombia; 10. C. palmeri (Riley & Howard)   from Argentina, Chile and Mexico; 11. C. pimentae (Newstead)   from Jamaica and USA. 12. C. rileyi (Cockerell) from Mexico; 13. C. rosae (Riley & Howard)   from Colombia, Mexico, Puerto Rico and USA (Florida); 14. C. similis (Morrison)   from El Salvador, Panama and Trinidad; 15. C. subandina (Leonardi)   from Argentina, and 16. C. zeteki (Cockerell)   from Colombia and Panama Canal Zone. Echinicerya   includes only E. anomala Morrison   from Guatemala. The most widespread monophlebid species in the Neotropics is Icerya purchasi Maskell   , found in most South American countries, but native to Australia. Two other Icerya   species have been recorded in the Neotropics, namely I. seychellarum (Westwood)   and I. schrottkyi Hempel.  

The tribe Llaveiini includes four genera: Laurencella   includes L. colombiana Foldi & Watson   from Colombia, L. marikana Foldi   from Mexico, L. taunayi (Hempel)   from Brazil and L. uhleri (Signoret)   (see below). Llaveia   includes L. axin (Llave)   from Guatemala, Mexico (Nearctic and Neotropical), and L. championi (Cockerell)   from Panama. Neohodgsonius   includes N. cassicola (Foldi)   from Venezuela. The species Palaeococcus brasiliensis (Walker)   was described based on adult males from Brazil ( Walker, 1852) and, although it is impossible to determine which genus this species belongs to, almost certainly it is not congeneric with the type species of Palaeococcus Cockerell   from Europe. In addition, Williams & Gullan (2008) have revised the Neotropical genus Protortonia Townsend   , describing and illustrating all known species, including all available instars, and this genus includes: 1. P. azteca (Ferris)   from Mexico; 2. P. cacti   (L.) from Puerto Rico; 3. P. c ro t o n i s Reyne from Netherlands Antilles; 4. P. ecuadorensis Foldi   from Ecuador and Colombia. 5. P. navesi Fonseca   from Brazil. 6. P. primitiva (Townsend)   from Mexico, and 7. P. quernea Williams & Gullan   from Costa Rica, collected at 3000 m on Quercus   sp. ( Fagaceae   ), on the slope of a volcano. Williams & Gullan (2008) provided keys to the genera in the tribe Llaveiini (based on adult female characters), and to the adult females of Protortonia   species. They transferred Marchalina azteca Ferris   to Protortonia   (as P. azteca (Ferris))   , synonymised Ecuadortonia Ben-Dov   with Laurencella Foldi   , and transferred Ortonia uhleri Signoret   to Laurencella   (as L. uhleri (Signoret))   .

Remark. Readers can refer to the key in Unruh & Gullan (2008 b) for identification of adult females of Iceryini and to Williams & Gullan (2008) for genera of Llaveiini.