Valid Names Results

Icerya Signoret, 1876 (Monophlebidae)

Nomenclatural History

• Icerya Signoret 1876: 351. Type species: Coccus sacchari Guerin-Meneville, 1867 by monotypy . (= Dorthesia seychellarum, Westwood, 1855 ) accepted valid name
• Crossotosoma Douglas 1890: 79. . junior synonym (discovered by Morris1928: 219)
• Iceryas Herrera 1903: 445;. . misspelling of genus name
• Iceria Hempel 1912: 57;. . misspelling of genus name
• Newsteadiella MacGillivray 1921: 75. Type species: Icerya formicarum Newstead by monotypy . junior synonym (discovered by Rao1951: 40)
• Auloicerya Morrison, H. {in}: Morrison & Morrison 1923: 22. Type species: Palaeococcus australis (Maskell) by original designation . junior synonym (discovered by UnruhGu2008: 41) Notes: The footnote on page 22 of Morrison & Morrison (1923) states "New genera should be credited to the senior author only", therefore the author of Auloicerya is Morrison, H.
• Aulo cerya Vayssière 1926: 306;. . misspelling of genus name
• Iceria Kiritchenko 1932a: 253;. . misspelling of genus name
• Pericerya Silvestri 1939: 648. Type species: Icerya purchasi Maskell by monotypy . junior synonym (discovered by MorrisMo1966: 152)
• Acerya Rao 1943: 246;. . misspelling of genus name
• Iceria Liegeois 1944: 165;. . misspelling of genus name
• Aulycerya Castel-Branco 1952: 23;. . misspelling of genus name
• Iceria Kaussari 1957: 2;. . misspelling of genus name
• Perycerya Gomez-Menor Ortega 1957: 86;. . misspelling of genus name
• Iceria Kawecki 1965a: 206;. . misspelling of genus name
• Perycerya Gomez-Menor Ortega 1968: 563;. by present designation . misspelling of genus name
• Icerva Hadzibejli 1983: 55;. . misspelling of genus name
• Preicerya Hadzibejli 1983: 55;. . misspelling of genus name
• Ycerya Hadzibejli 1983: 266;. . misspelling of genus name
• Auloicerya Unruh & Gullan 2008: 41. . subsequent use Notes: The authorships of Auloicerya is incorrectly attributed to Morrison & Morrison.

Remarks

• Systematics: Main characteristics (adult female) of this genus are: derm dorsally and marginally strongly sclerotized; 10- or 11-segmented antenna; labium conical, one-segmented; legs well-developed; three pairs of abdominal spiracles; with ventral abdominal cicatrices.

  Unruh & Gullan (2008b) recognised 8 species groups as follows:

  Icerya aegyptiaca group Two species belong to this group: I. aegyptiaca and I. schrottkyi. The pores of these two species are quite different, but both have pores on the derm that resemble the vulvar pores. They are separated easily by the number of cicatrices (I. aegyptiaca has one, I. schrottkyi has three) and are separated further by the shape of pores on the derm. Icerya schrottkyi has unique hexalocular pores on the ventral surface that are absent from all other iceryine species.

  Icerya jacobsoni group Unruh & Gullan (2008b) included five species in this group: I. assamensis, I. jacobsoni, I. jaihind, I. mangiferae and I. zimmermanni. Although the species vary in their egg-tending behaviour I. assamensis forms an internal marsupium, I. zimmermanni has an ovisac band and a simple waxy pad of secretion covering the ventral surface and I. jacobsoni, I. jaihind and I. mangiferae form neither an ovisac nor a marsupium), simple multilocular pores with characteristic stalked profiles are found on the dorsal surface of all species. Icerya jacobsoni and I. jaihind differ from one another by the number of ventral cicatrices (I. jacobsoni has three and I. jaihind has a single cicatrix). Unruh & Gullan (2008b) were unable to examine material of I. mangiferae, but included it in this group based on the original description and because they believed it a likely synonym of I. jacobsoni. Refer to the taxonomic notes on I. mangiferae for further discussion.

  Icerya natalensis group Three species make up this group: I. clauseni, I. natalensis and I. travancorensis. The former two species are the only Icerya species that do not form an ovisac band and have open-centre pores. The open-centre pores of each species resemble each other except that the open-centre pores of I. clauseni have a small cleft, which is lacking from the open-centre pores of I. natalensis. The third species, I. travancorensis lacks open-centre pores and forms an ovisac band. All species have only simple multilocular pores with bilocular centres on the dorsal surface, but I. natalensis differs by the shape of the pores on the ventromedial abdomen [I. natalensis has larger pores that appear bluish when stained] with a trilocular centre and 8-12 elongate outer loculi, while I. clauseni and I. travancorensis have smaller pores with a bilocular or trilocular centre and 4-6 outer loculi]. Icerya natalensis is very rare and was collected originally in Natal, South Africa, over 100 years ago. Unruh & Gullan (2008b) placed it in Icerya rather than Gigantococcus, because of the presence of open-centre pores and absence of compound multilocular pores.

  Icerya nudata group Unruh & Gullan (2008b) included two marsupium-forming species in this group: I. nudata and I. samaraia. These are the only two marsupium-forming species that have open-centre pores present on the derm. They can be separad by the shape and size of the open-centre pores: I. samaraia has large pores with tightly-spaced outer loculi while the open-centre pores of I. nudata are smaller, have fewer widely-spaced outer loculi and have a small cleft.

  Icerya pilosa group Unruh & Gullan (2008b) placed two species in this group: Icerya pilosa and I. kumari. Unruh & Gullan (2008b) were unable to examine material of I. kumari. Based on Rao's description, illustration and photographs, they believed it was probably closely related to Icerya pilosa and that it might even be a third-instar nymph of that species (refer to the taxonomic notes on I. kumari for further discussion). On both species, the derm around the open-centre pores becomes very sclerotized and the loculi present in the outer rim look similar. Icerya pilosa forms an ovisac band but Rao believed I. kumari formed neither an ovisac band nor a marsupium.

  Icerya pulchra group This group includes three species: I. pulchra, I. morrisoni and I. minor. All have the same pore types distributed in the same way on the derm. Icerya morrisoni has a single cicatrix whereas I. minor and I. pulchra have three cicatrices. The latter two species differ by the density of simple multilocular pores with a quinquelocular or hexalocular centre and 4-8 outer loculi. Morrison believed I. minor and I. pulchra to be synonyms because the only difference he found between the two species was the number of multilocular pores on the dorsum. Green also believed these two species to be synonyms and suggested that I. minor represented a smaller form of I. pulchra. Rao separated the two species based on the length of the antennal and apical setae of the first-instar nymph (long in I. minor, short in I. pulchra). He found that the dorsal pores of I. minor tend to have bilocular centres while the dorsal pores of I. pulchra tend to have trilocular centres.

  Icerya seychellarum group Five species belong to this group: I. crocea, I. formicarum, I. hanoiensis, I. menoni and I. seychellarum. Until recently, I. crocea was considered a synonym of I. seychellarum, but the two were separated based on the shape and distribution of the open-centre pores and genetic differences (Unruh & Gullan, 2008). These five species all have open-centre pores present in marginal clusters and transverse rows on the dorsal surface. This distribution of open-centre pores differs from other species in which the distribution is restricted to the margins and across the head only. The five species of this group can be separated based on the shape and density of their open-centre pores. The open-centre pores of I. crocea have 17-20 outer loculi and may or may not have a small cleft present. These pores are densely clustered on the middorsal head and thorax. Icerya seychellarum has similar-looking pores that always have a triangular cleft and are scattered only across the dorsal head and thorax. Both species have three cicatrices, which differs from I. formicarum and I. menoni, which each have a single cicatrix. The latter two species can be separated by the number of loculi in the open-centre pores and the shape of the ovisac in life [we were unable to examine material of I. menoni and base these claims on Rao's (1951a) description and illustrations]. In life, the external appearance of the two species differs as the ovisac of I. menoni consists of a fluffy white secretion on the ventral abdomen compared to the elongate, fluted ovisac of I. formicarum. Icerya menoni also has two pencils of wax projecting from the posterior end of the body [projections visible in photograph accompanying original description (Rao, 1951a: 61)] and I. formicarum has two tufts of wax surrounding the anal opening (1951a: 54). Rao reported that the open-centre pores of I. menoni are larger than those of I. formicarum and have a proportionately larger number of outer loculi and larger diameter. Refer to the taxonomic notes (in Unruh & Gullan, 2008b) on I. hanoiensis for a discussion of possible synonymy with I. seychellarum.

  "Pericerya" group Species in this group form a clade in the molecular phylogenetic analysis by Unruh & Gullan, 2008. "Pericerya" is the generic name created by Silvestri (1939) to describe all iceryine species with two pairs of abdominal spiracles. Four species, I. purchasi, I. callitri, I. koebelei and I. sumatrana, as well as several undescribed species from New Caledonia and Australia, belong to the "Pericerya" group. All species have only two pairs of abdominal spiracles and have open-centre pores present in marginal clusters. Icerya callitri looks the most dissimilar to the other species with simple multilocular pores, similar to vulvar pores, present on the dorsal surface and ventral margin to submargin. Icerya purchasi and I. koebelei can be separated easily in life because I. koebelei forms a "pencil" of wax that projects upward from the middorsum. Under the microscope they can be distinguished from one another by the number of antennal segments (I. purchasi has 11 segments, I. koebelei typically has 9 or 10) and by the density of dark hair-like setae on the dorsal surface. Also I. koebelei has a dense middorsal cluster of 30-50 simple multilocular pores from which the wax pencil emanates. Icerya sumatrana resembles I. purchasi in life, but can be separated by the shape of the multilocular pores, the absence of marginal clusters of setae and the absence of bilocular centre pores on the dorsum.

  Unplaced Icerya species Four Icerya species that were examined by Unruh & Gullan (2008b) could not be placed in any other group. These species are I. australis, I. acaciae, I. imperatae and I. nuda. Unruh & Gullan (2008b) did not place I. nuda in any group because the material was in very poor shape. Several undescribed species of Icerya that look similar to I. imperatae have been collected in Australia. Although I. acaciae and I. australis look similar and previously belonged to the genus Auloicerya, they were not closely related in the phylogenetic reconstruction of Unruh & Gullan (2008b).

  Gavrilov-Zimin, 2018, resurrected the tribe Iceryini within the family Margarodidae to place the genus Icerya in relation to other genera that he felt belonged in a subfamily of Monophlebinae (Crypticerya , Echinicerya, Icerya and Steatococcus) He rejected the molecular evidence for the separation of genera in favor of a phylogenetic evaluation of reproductivde strategies in different "traditionally accepted genera." Based on his morphological definition of the separation of genera, he considered Steatococcus a separate genus and reassigned several species in Crypticerya to the genus Steatococcus.
• General Remarks: Definition and description by Morrison & Morrison (1923), Vayssiere (1926), Morrison (1928), Unruh & Gullan (2008) and by Unruh & Gullan (2008b).

Keys

• Watson2022: pp.320-321 ( Adult (F) ) [Monophlebidae in continental Africa]
• UesatoKoUn2011: pp.52 ( Adult (F) ) [Icerya species in Japan]
• UnruhGu2008: pp.23-24 ( Adult (F) ) [Genera of Margarodidae - Iceryini]
• UnruhGu2008: pp.24 ( First instar ) [Genera of Margarodidae - Iceryini]
• UnruhGu2008b: pp.22-23 ( Adult (F) ) [Genera of Iceryini]
• UnruhGu2008b: pp.72-73 ( Adult (F) ) [Species of Icerya]
• Koszta1996: pp.40 ( Adult (F) ) [North America]
• Gill1993: pp.35 ( Adult (F) ) [USA, California]
• Morale1991: pp.56 ( Adult (F) ) [New Zealand]
• WilliaWa1990: pp.15-16 ( Adult (F) ) [Margarodidae Genera of Tropical South pacific]
• WilliaWa1990: pp.21 ( Adult (F) ) [Icerya species of South Pacific]
• Beards1966: pp.391 ( Adult (F) ) [Genera of Micronesia]
• Borchs1950b: pp.32 ( Adult (F) ) [Genera of USSR]
• Morris1928: pp.196-197 ( Immature (F) ) [Margarodidae]
• Vayssi1926: pp.258-259 ( Adult (F) ) [Margarodidae]
• Vayssi1923a: pp.428-429 ( Adult (F) ) [Monophlebinae]
• Kuwana1922a: pp.2 ( Adult (F) ) [Japan]
• MacGil1921: pp.73-74 ( Adult (F) ) [Genus]
• Morris1920: pp.150 ( Adult (F) ) [Philippines]
• RileyHo1890b: pp.106 ( Immature (F) Immature (M) ) [USA, Florida]

Associated References

• Beards1966: taxonomy, pp. 397
• BenDov2005a: catalog, taxonomy, pp. 150,186-188
• Bodenh1953a: taxonomy, pp. 155
• Borchs1950b: taxonomy, pp. 32, 45
• Brain1915: taxonomy, pp. 166-167
• Cocker1895z: description, taxonomy, pp. 15
• Cocker1899m: taxonomy, pp. 274
• Cocker1899n: taxonomy, pp. 3, 4
• Cocker1902o: taxonomy, pp. 80
• Cocker1902q: taxonomy, pp. 257
• Comsto1881a: taxonomy, pp. 347
• Dougla1890: taxonomy, pp. 79
• Fernal1903b: pp. 23
• Ferris1919a: description, taxonomy, pp. 11-12
• Foldi1998: catalog, pp. 424-425
• Foldi2001a: taxonomy, pp. 197,202
• Frogga1921b: taxonomy, pp. 37
• Fuller1897b: taxonomy, pp. 1346
• Fuller1897c: taxonomy, pp. 11
• Fuller1899: taxonomy, pp. 438
• Gill1993: description, taxonomy, pp. 35, 36
• GomezM1937: description, taxonomy, pp. 395
• GomezM1957: taxonomy, pp. 86
• GomezM1968: taxonomy, pp. 563
• Green1922: description, taxonomy, pp. 435-436
• GullanSj2001: taxonomy, pp. 257-278
• Hadzib1983: taxonomy, pp. 55, 266
• Hempel1900a: taxonomy, pp. 370
• Hempel1912: taxonomy, pp. 17, 57
• HodgsoFo2006: taxonomy, pp. 162
• HodgsoLa2011: taxonomy, pp. 2
• Kaussa1957: taxonomy, pp. 2
• Kawai1980: description, taxonomy, pp. 87
• Kaweck1965a: taxonomy, pp. 206
• Kiritc1932a: taxonomy, pp. 253
• Koszta1996: description, taxonomy, pp. 40-41
• Kuwana1922a: description, taxonomy, pp. 1-2
• Liegeo1944: taxonomy, pp. 165
• Lindin1937: taxonomy, pp. 182, 193
• Lindin1943b: taxonomy, pp. 221, 222
• Lizery1939: catalog, taxonomy, pp. 172
• MacGil1921: taxonomy, pp. 71,73-75
• Mamet1949: catalog, taxonomy, pp. 4
• Maskel1887a: taxonomy, pp. 90
• MkrtchSa2001: structure, taxonomy, pp. 167-168
• Morale1991: description, taxonomy, pp. 56
• Morris1920: taxonomy, pp. 149-150
• Morris1928: description, taxonomy, pp. 6-26,100-220
• MorrisMo1923: pp. 22-28,31
• MorrisMo1966: taxonomy, pp. 48, 95, 134, 152, 16
• Rao1943VP: taxonomy, pp. 246
• Rao1951: description, taxonomy, pp. 40-50
• RileyHo1890b: taxonomy, pp. 92-106
• Signor1876: description, taxonomy, pp. 351-352
• Silves1920: taxonomy, pp. 487
• SinghKa2017: host, pp. 258
• TangHa1995: description, taxonomy, pp. 147, 154-156
• Tao1999: taxonomy, pp. 2-3
• UesatoKoUn2011: taxonomy, pp. 49-55
• UnruhGu2005: taxonomy, pp. 133
• UnruhGu2008: description, molecular data, phylogeny, taxonomy, pp. 8-50
• UnruhGu2008b: description, taxonomy, pp. 69-73
• Varshn2005: taxonomy, pp. 142
• Vayssi1923a: description, taxonomy, pp. 428-429
• Vayssi1926: description, taxonomy, pp. 259,306-307,316
• Watson2022: key, pp. 320
• WilliaWa1990: description, taxonomy, pp. 16,18-21
• Yang1982: taxonomy, pp. 19, 30-31
• Zahrad1959a: taxonomy, pp. 534-535
• Zimmer1948: taxonomy, pp. 136

38 Species