Coccoidea

Female and male of Apiomorpha, copyright Galler18.

Belongs within: Coccinea.
Contains: Lecanodiaspididae, Asterolecaniidae, Tachardiidae, Pseudococcidae, Eriococcidae, Coccidae, Diaspididae, Puto.

The Coccoidea are recognised here, following Koteja (2000), as including the neococcoid scale insects; other authors use this name to refer to all scale insects (Coccinea). The earliest representatives are known from Upper Cretaceous amber, with the extinct taxa Inka minuta and Labiococcidae known only from this period (Koteja 2000). Modern representatives include the Kerriidae, a primarily tropical group with brachial plates and a dorsal spine (Carver et al. 1991). Resinous secretions from the lac insect Kerria lacca are used to produce shellac.

Characters (from Grimaldi & Engel 2005, as 'neococcoids'): Females lacking abdominal spiracles; male eyes reduced to individual facets; unique chromosomal system of sex determination by paternal genome elimination.

<==Coccoidea K00
    |--+--Pseudococcidae GE05
    |  `--+--Inka Koteja 1989 K00 [Inkaidae GE05]
    |     |    `--I. minuta Koteja 1989 K00
    |     `--+--Eriococcidae GE05
    |        |--Coccidae GE05
    |        `--Diaspididae GE05
    `--+--Puto GE05
       `--Labiococcidae GE05
            |--Labiococcus Koteja 2000 K00
            |    `--L. joosti Koteja 2000 K00
            `--Solicoccus Koteja 2000 K00
                 `--S. nascimbenei Koteja 2000 K00

Coccoidea incertae sedis:
  Kermesidae K00
    |--Kermes nigropunctatus M96
    |--Olliffiella secunda M96
    |--Sucinikermes Koteja 1988 K00
    |    `--S. kulickae Koteja 1988 K00
    `--Allokermes M96
         |--A. galliformis M96
         `--A. grandis M96
  Aclerda [Aclerdidae] WEE70
    |--A. arundinariae S96
    |--A. attenuata M96
    |--A. campinensis S96
    |--A. manni M96
    |--A. mexicana M96
    |--A. oaxacaensis M96
    |--A. sellahispanica WEE70
    |--A. sinaloaensis M96
    `--A. takahashii WEE70
  Lecanodiaspididae K00
  Beesoniidae K00
  Micrococcidae K00
  Dactylopius TRZ10 [Dactylopiidae K00]
    |--D. ceylonicus B82
    |--D. coccus TRZ10
    |--D. confusus G10
    |--D. glaciale G17
    |--D. kingii C01
    |--D. neomexicanus C01
    |--D. opuntiae CGW91
    `--D. tomentosus M96
  Apiomorpha CGW91 [Apiomorphidae K00]
    |--A. conica G84
    |--A. duplex AY04
    |--A. frenchi CGW91
    |--A. maliformis B88
    |--A. munita CGW91
    |--A. ovicola [=Brachyscelis ovicola] B88
    |--A. pedunculata N91
    |--A. pileata [=Brachyscelis pileata] B88
    `--A. strombylosa [incl. Brachyscelis crispa] B88
  Cerococcus CGW91 [Cerococcidae K00]
    |--C. koebelei M96
    |--C. quercus RD77
    |--C. russellae M96
    |--C. stellatus CGW91
    `--C. zapotlanus M96
  Colobopyga CGW91 [Halimococcidae GE05]
    |--C. attaleae M96
    |--C. australiensis CGW91
    |--C. coperniciae WEE70
    |--C. kewensis CGW91
    |--C. sabalis M96
    `--C. washingtoniae M96
  Phoenicococcus [Phoenicococcidae] T02
    `--P. marlatti CK15
  Stictococcus G17 [Stictococcidae CGW91]
    `--S. gowdeyi G17
  Asterolecaniidae CGW91
  Kerriidae [Lacciferidae] CGW91
    |--Kerria lacca GE05
    |--Austrotachardia angulata CGW91
    |--Tachardina CGW91
    |--Laccifer lacca WEE70
    `--Gascardia destructor RD77, DAF07
  Conchaspis [Conchaspididae] CGW91
    |--C. angraeci CGW91
    `--C. newsteadi M96
  Tachardiidae M96
  Cryptococcus fagi RD77
  Physokermes RD77
    |--P. insignicola G17
    `--P. piceae RD77
  Parthenolecanium corni RD77

*Type species of generic name indicated

REFERENCES

[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. pHales & G. S. Taylor. 2004. Insects 'Down Under'—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216–234.

[B82] Batra, S. W. T. 1982. Biological control in agroecosystems. Science 215: 134–139.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[CK15] Calvo, F. J., M. Knapp, Y. M. van Houten, H. Hoogerbrugge & J. E. Belda. 2015. Amblyseius swirskii: what made this predatory mite such a successful biocontrol agent? Experimental and Applied Acarology 65: 419–433.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[C01] Cockerell, T. D. A. 1901. Contributions from the New Mexico Biological Station.—XI. New and little-known insects from New Mexico. Annals and Magazine of Natural History, series 7, 7: 333–337.

[DAF07] Department of Agriculture and Food. 2007. Common Seasonal Pests. State of Western Australia.

[G10] Gaimari, S. D. 2010. Chamaemyiidae (chamaemyiid flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 997–1007. NRC Research Press: Ottawa.

[G84] Gauld, I. D. 1984. An Introduction to the Ichneumonidae of Australia. British Museum (Natural History).

[G17] Girault, A. A. 1917. Descriptiones stellarum novarum. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 80–101).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[K00] Koteja, J. 2000. Advances in the study of fossil coccids (Hemiptera: Coccinea). Polskie Pismo Entomologiczne 69: 187–218.

[M96] Miller, D. R. 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington 98 (1): 68–86.

[N91] Norris, K. R. 1991. General biology. In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 68–108. Melbourne University Press: Carlton (Victoria).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S96] Sharkov, A. 1996. A review of the species of the genus Mucrencyrtus Noyes (Hymenoptera: Encyrtidae). Proceedings of the Entomological Society of Washington 98 (2): 350–368.

[T02] Takagi, S. 2002. One new subfamily and two new tribes of the Diaspididae (Homoptera: Coccoidea). Insecta Matsumurana, new series 59: 55–100.

[TRZ10] Thompson, E. C., G. E. Rotheray & M. A. Zumbado. 2010. Syrphidae (flower flies). In: Brown, B. V., A. Borkent, J. M. Cumming, D. M. Wood, N. E. Woodley & M. A. Zumbado (eds) Manual of Central American Diptera vol. 2 pp. 763–792. NRC Research Press: Ottawa.

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

Last updated: 1 May 2022.

Coccinea

Cottony cushion scale Icerya purchasi, from Clemson University.

Belongs within: Sternorrhyncha.
Contains: Ortheziidae, Coccoidea, Matsucoccidae.

The Coccinea, scale insects, are a group of plant-sucking bugs in which the females have become larviform and generally immobile. Males are winged and non-feeding at maturity; the hind pair of wings is reduced and may be represented only by a pair of bristle-like structures. First-instar nymphs of both sexes (known as 'crawlers') are highly mobile, dispersing in search of suitable host plants before moulting into the subsequent immobile stage.

Characters (from Griamldi & Engel 2005, as Coccoidea): Sexual dimorphism extreme: females highly neotenic, larviform, lacking eyes and wings, antennae and legs also usually highly absent; first-instar crawler as dispersal stage. Males lacking mouthparts; hind pair of wings reduced, often to tiny knobs (hamulohalteres).

<==Coccinea [Coccidomorpha, Coccina, Coccomorpha, Orthezioidea] KP01
    |  i. s.: Polyclona Menge 1856 K00
    |         Neomargarodes erythrocephala K00
    |         Permaleurodes GE05
    |         Carayonemidae GE05
    |         Ortheziola vejdovskyi K09
    |         Lophococcus maximus F09
    |--Mesococcus K00
    |    |--M. asiaticus Becker-Migdisova in Rodendorf 1962 K00
    |    `--M. lutarius Zhang 1985 RJ93
    `--+--Ortheziidae GE05
       `--+--Coccoidea GE05
          |--Kukaspis Koteja & Poinar 2001 KP01 [Kukaspididae GE05]
          |    `--*K. usingeri Koteja & Poinar 2001 KP01
          |--Steingelia Nassonov 1908 K00 [Steingeliidae GE05]
          |    `--S. cretacea Koteja 2000 K00
          |--Phenacoleachia [Phenacoleachiidae] GE05
          |    `--P. zealandica GE05
          `--+--Matsucoccidae GE05
             `--+--+--Cancerococcus Koteja 1988 K00 [Pityococcidae GE05]
                |  |    `--C. apterus Koteja 1988 K00
                |  `--Electrococcidae GE05
                |       |--Electrococcus Beardsley 1969 K00
                |       |    `--E. canadensis Beardsley 1969 K00
                |       `--Turonicoccus Koteja 2000 K00
                |            |--T. beardsleyi Koteja 2000 K00
                |            `--T. grimaldii Koteja 2000 K00
                `--+--+--Coelostomidiidae [Coelostomidiinae] GE05
                   |  |    |--Coelostomidia zealandica WD04
                   |  |    `--Ultracoelostoma WD04
                   |  |         |--U. assimile WD04
                   |  |         `--U. brittini WD04
                   |  `--+--Jersicoccus Koteja 2000 K00 [Jersicoccidae GE05]
                   |     |    `--J. kurthi Koteja 2000 K00
                   |     `--Monophlebus K00 [Monophlebidae GE05]
                   |          |--M. irregularis Germar & Berendt 1856 K00
                   |          `--M. simplex Scudder 1890 K00
                   `--+--Baisococcus Koteja 1989 K00 [Xyloccidae, Xylococcidae GE05]
                      |    `--B. victoriae Koteja 1989 K00
                      `--+--Grimaldiella Koteja 2000 K00 [Grimaldiellidae GE05]
                         |    |--G. gregaria Koteja 2000 K00
                         |    `--G. resinophila Koteja 2000 K00
                         `--Margarodidae GE05
                              |--Melaleucococcus nodosus GE05
                              |--Callipappus rubiginosus GE05, B88
                              |--Monophlebulus pilosior S07
                              |--Auloicerya CGW91
                              |--Nodulicoccus CGW91
                              |--Conifericoccus agathidis CGW91
                              |--Eumargarodes laingi CGW91
                              |--Promargarodes australis CGW91
                              |--Icerya [Iceryini] CGW91
                              |    |--I. aegyptiaca F04
                              |    |--I. colimensis M96
                              |    |--I. littoralis M96
                              |    |    |--I. l. littoralis M96
                              |    |    |--I. l. mimosae M96
                              |    |    `--I. l. tonilensis M96
                              |    |--I. montserratensis M96
                              |    |--I. palmeri M96
                              |    |--I. purchasi S07
                              |    |--I. rileyi M96
                              |    |    |--I. r. rileyi M96
                              |    |    `--I. r. larreae M96
                              |    `--I. seychellarum B88
                              |--Cryptokermes Hempel 1901 M96, H01
                              |    |--*C. brasiliensis Hempel 1901 H01
                              |    `--C. mexicanus M96
                              |--Llaveia M96
                              |    |--L. axin M96
                              |    |--L. mexicanorum M96
                              |    `--L. oaxacoensis M96
                              |--Marchalina azteca M96
                              |--Neosteingelia texana M96
                              |--Protortonia primitiva M96
                              |--Steatococcus M96
                              |    |--S. mexicanus M96
                              |    |--S. morrilli M96
                              |    `--S. tabernicolus M96
                              `--Margarodes hiemalis W87

*Type species of generic name indicated

REFERENCES

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[F04] Ferrar, P. 2004. Australian entomology: isolated, or in touch with the rest of the world? Australian Journal of Entomology 43 (3): 329–333.

[F09] Froggatt, W. W. 1909. Notes and exhibits. Proceedings of the Linnean Society of New South Wales 33: 798.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H01] Hempel, A. 1901. Descriptions of Brazilian Coccidae. Annals and Magazine of Natural History, series 7, 7: 110–125.

[K09] Köhler, G. 2009. Heuschrecken (Saltatoria) und Ohrwürmner (Dermaptera) im Immissionsgebiet des Düngemittelwerkes Steudnitz/Thüringen—eine Langzeitstudie (1978–2001). Mauritiana 20 (3): 601–646.

[K00] Koteja, J. 2000. Advances in the study of fossil coccids (Hemiptera: Coccinea). Polskie Pismo Entomologiczne 69: 187–218.

[KP01] Koteja, J., & G. O. Poinar Jr. 2001. A new family, genus, and species of scale insect (Hemiptera: Coccinea: Kukaspididae, new family) from Cretaceous Alaskan amber. Proceedings of the Entomological Society of Washington 103 (2): 356–363.

[M96] Miller, D. R. 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington 98 (1): 68–86.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[S07] Ślipiński, A. 2007. Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their biology and classification. Australian Biological Resources Study: Canberra.

[W87] Walter, D. E. 1987. Belowground arthropods of semiarid grasslands. In: J. L. Capinera (ed.) Integrated Pest Management on Rangeland: A shortgrass prairie perspective pp. 271–290. Westview Press, Inc.: Boulder (Colorado).

[WD04] Wardhaugh, C. W., & R. K. Didham. 2004. The effect of introduced wasp (Vespula vulgaris, Hymenoptera: Vespidae) predation on the dispersive life history stages of beech scale insects (Ultracoelostoma spp., Homoptera: Margarodidae). New Zealand Entomologist 27: 91–101.

Last updated: 2 April 2022.

Eriococcidae

Gum-tree scale Acanthococcus coriaceus, photographed by Arthur Chapman.

Belongs within: Coccoidea.

The Eriococcidae are a group of scale insects characterised by well-developed anal lobes, a poriferous and setiferous anal ring, conical or bulbous dorsal setae, enlarged macroducts cupped at the inner end, microducts, quinquelocular disc pores, and small cruciform pores. Representatives sometimes cause remarkable galls on host trees (Carver et al. 1991). Some are significant pests on tree species such as Acanthococcus coriaceus on Eucalyptus or A. ironsidei on macadamia.

Characters (from Carver et al. 1991): Female without dorsal ostioles, ventral circuli and cerarii; anterior and posterior pairs of thoracic spiracles approximately the same size; abdominal spiracles absent; terminal segments of abdomen not fused into pygidium; anal ring usually with a single row of pores; tubular ducts, if present, with inner end invaginated to form a cup.

<==Eriococcidae [Idiococcinae]
    |--Madarococcus MC94
    |--Opisthoscelis AY04
    |--Cystococcus CGW91
    |--Lachnodius eucalypti B88
    |--Epicoccus CGW91
    |--Sphaerococcopsis CGW91
    |--Ascelis CGW91
    |--Casuarinaloma CGW91
    |--Floracoccus CGW91
    |--Gossyparia casuarinae WEE70, S07
    |--Olliffia WEE70
    |--Ourococcus WEE70
    |--Capulinia sallei M96
    |--Carpochloroides mexicanus M96
    |--Oregmopyga peninsularis M96
    |--Xerococcus fouquieriae M96
    |--Kaweckia MG96
    |--Apezococcus idiastes W87
    |--Keithia Koteja 2000 nec Spreng. 1822 (ICBN) nec Sacc. 1892 (ICBN) K00
    |    `--K. luzzii Koteja 2000 K00
    |--Gedanicoccus Koteja 1988 K00
    |    `--G. gracilis Koteja 1988 K00
    |--Kuenowicoccus Koteja 1988 K00
    |    `--K. pietrzeniukae Koteja 1988 K00
    |--Balticoccus Koteja 1988 K00
    |    |--B. oblicus Koteja 1988 K00
    |    `--B. spinosus Koteja 1988 K00
    |--Jutlandicoccus Koteja 1988 K00
    |    |--J. pauper Koteja 1988 K00
    |    `--J. perfectus Koteja 1988 K00
    |--Eriococcus CGW91
    |    |--E. nitidus B88
    |    `--E. rorismarinus Fonscolombe 1834 S69
    |--Cylindrococcus [Cylindrococcidae] N91
    |    |--C. amplior G34
    |    |--C. casuarinae B88
    |    `--C. spiniferus CGW91
    `--Acanthococcus Signoret 1875 [incl. Gregoporia Danzig 1979] MG96
         |--A. abdifus (Hoy 1962) [=Eriococcus abditus] MG96
         |--A. acericola (Tang & Hao 1995) [=Eriococcus acericola] MG96
         |--A. aceris MG96
         |    |--A. a. aceris MG96
         |    `--A. a. kurdicus (Bodenheimer 1943) [=Eriococcus aceris kurdica] MG96
         |--A. acutispinatus (Hoy 1962) [=Eriococcus acutispinatus] MG96
         |--A. agonis (Fuller 1897) [=Eriococcus agonis] MG96
         |--A. albatus (Hoy 1962) [=Eriococcus albatus] MG96
         |--A. amomidis (Gomez-Menor 1935) [=Eriococcus amomidis] MG96
         |--A. angulatus (Froggatt 1916) [=Eriococcus angulatus] MG96
         |--A. apiomorphae (Fuller 1897) [=Eriococcus apiomorphae] MG96
         |--A. araucariae M96
         |    |--A. a. araucariae MG96
         |    |--A. a. minor (Maskell 1895) [=Eriococcus araucariae minor] MG96
         |    `--A. a. nudus (Gomez-Menor 1957) [=Eriococcus araucariae nuda] MG96
         |--A. arboisi (Goux 1991) [=Eriococcus arboisi] MG96
         |--A. arcanus (Hoy 1962) [=Eriococcus arcanus] MG96
         |--A. argentifagi (Hoy 1962) [=Eriococcus argentifagi] MG96
         |--A. armeniacus (Tang & Hao 1995) [=Eriococcus armeniacus] MG96
         |--A. artiguesi (Goux 1991) [=Eriococcus artiguesi] MG96
         |--A. asteliae (Hoy 1962) [=Eriococcus asteliae] MG96
         |--A. aurescens M96
         |--A. australis (Maskell 1895) [=Eriococcus buxi australis] MG96
         |--A. bambusae (Green 1922) [=Eriococcus bambusae] MG96
         |--A. beilschmiediae (Hoy 1962) [=Eriococcus beilschmiediae] MG96
         |--A. betulaefoliae (Tang & Hao 1995) [=Eriococcus betulaefoliae] MG96
         |--A. bezzii (Leonardi 1907) [=Eriococcus bezzii] MG96
         |--A. bicolor (Froggatt 1915) [=Rhizococcus bicolor] MG96
         |--A. boguschi (McDaniel 1964) [=Eriococcus boguschi] MG96
         |--A. brachypodii (Borchsenius & Danzig 1966) [=Greenisca brachypodii] MG96
         |--A. brasiliensis (Cockerell 1900) [=Eriococcus brasiliensis] MG96
         |--A. brevenniae (Goux 1993) [=Eriococcus brevenniae] MG96
         |--A. brittini (Hoy 1962) [=Eriococcus brittini] MG96
         |--A. cactearum (Leonardi 1918) [=Eriococcus cactearum] MG96
         |--A. campbelli (Hoy 1959) [=Eriococcus campbelli] MG96
         |--A. campinensis (Hempel 1937) [=Eriococcus campinensis] MG96
         |--A. cantium (Williams 1985) [=Eriococcus cantium] MG96
         |--A. castanopus (Tang & Hao 1995) [=Eriococcus castanopus] MG96
         |--A. casuarinae (Maskell 1893) [=Rhizococcus casuarinae] MG96
         |--A. caudatus (Tang & Hao 1995) [=Eriococcus caudatus] MG96
         |--A. cavellii (Maskell 1890) [=Gossyparia cavellii] MG96
         |--A. celmisiae (Masekll 1884) [=Rhizococcus celmisiae] MG96
         |--A. chaoticus (Goux 1991) [=Eriococcus chaoticus] MG96
         |--A. chathamensis (Hoy 1962) [=Eriococcus chathamensis] MG96
         |--A. chilensis (Miller & González 1975) [=Eriococcus chilensis] MG96
         |--A. coccineus M96
         |--A. coffeae (Hempel 1919) [=Eriococcus coffeae] MG96
         |--A. confluens (Maskell 1893) [=Gossyparia confluens] MG96
         |--A. confusus (Maskell 1892) [=Eriococcus confusus] MG96
         |--A. consperus (Masekll 1893) [=Eriococcus consperus] MG96
         |--A. constrictus (Froggatt 1933) [=Eriococcus constrictus] MG96
         |--A. coprosmae (Hoy 1962) [=Eriococcus coprosmae] MG96
         |--A. coriaceus (Maskell 1893) [=Eriococcus coriaceus] MG96
         |--A. corniculatus (Ferris 1950) [=Eriococcus corniculatus] MG96
         |--A. costaricensis (Cockerell & Robinson 1915) [=Eriococcus costaricensis] MG96
         |--A. crenilobatus (Hoy 1962) [=Eriococcus crenilobatus] MG96
         |--A. crispus (Fonscolombe 1834) [=Coccus crispus] MG96
         |--A. crofti (Froggatt 1916) [=Eriococcus crofti] MG96
         |--A. cultellus (Hoy 1959) [=Eriococcus cultellus] MG96
         |--A. curassavicus (Reyne 1964) [=Eriococcus curassavicus] MG96
         |--A. dacrydii (Hoy 1962) [=Eriococcus dacrydii] MG96
         |--A. danthoniae (Maskell 1891) [=Eriococcus danthoniae] MG96
         |--A. danzigi Miller & Gimpel 1996 [=A. confusus Danzing 1962 non Eriococcus confusus Maskell 1892] MG96
         |--A. detectus (Hoy 1962) [=Eriococcus detectus] MG96
         |--A. devoniensis [=Eriococcus devoniensis] RD77
         |--A. diverspinus (Leonardi 1911) [=Eriococcus diverspinus] MG96
         |--A. dubius M96
         |--A. dziedzickae Miller & Gimpel 1996 (see below ofr synonymy) MG96
         |--A. elaeocarpi (Hoy 1962) [=Eriococcus elaeocarpi] MG96
         |--A. elegans (Fuller 1897) [=Eriococcus elegans] MG96
         |--A. elytranthae (Hoy 1962) [=Eriococcus elytranthae] MG96
         |--A. emirnensis (Mamet 1954) [=Eriococcus emirnensis] MG96
         |--A. ericae (Signoret 1875) [=Eriococcus ericae] MG96
         |--A. etbaicus (De Lotto 1954) [=Eriococcus etbaicus] MG96
         |--A. eucalypti (Maskell 1892) [=Eriococcus eucalypti] MG96
         |--A. eurythrix (Miller & González 1975) [=Eriococcus eurythrix] MG96
         |--A. evelinae (Kozar 1983) [=Rhizococcus evelinae] MG96
         |--A. fagicorticis (Maskell 1892) [=Eriococcus fagicorticis] MG96
         |--A. festucarum (Lindinger 1932) (see below for synonymy) MG96
         |--A. formicicola (Newstead 1897) [=Eriococcus formicicola] MG96
         |--A. fossilis (Froggatt 1933) [=Eriococcus fossilis] MG96
         |--A. fossor (Maskell 1884) [=Rhizococcus fossor] MG96
         |--A. franceschinii (Balachowsky 1934) [=Eriococcus franceschinii] MG96
         |--A. fuligitectus (Hoy 1962) [=Eriococcus fuligitectus] MG96
         |--A. gassinus (Goux 1992) [=Eriococcus gassinus] MG96
         |--A. gaultheriae (Hoy 1962) [=Eriococcus gaultheriae] MG96
         |--A. gerbergeri M96
         |--A. gibbus (Hoy 1959) [=Eriococcus gibbus] MG96
         |--A. glyceriae (Green 1921) [=Eriococcus glyceriae] MG96
         |--A. gouxi (Balachowsky 1954) [=Anophococcus gouxi] MG96
         |--A. graminicola (Borchsenius 1949) [=Rhizococcus graminicola] MG96
         |--A. grandis (Maskell 1892) [=Rhizococcus grandis] MG96
         |    |--A. g. grandis MG96
         |    `--A. g. spinosior (Maskell 1893) [=Rhizococcus grandis spinosior] MG96
         |--A. guesinus (Goux 1992) [=Eriococcus guesinus] MG96
         |--A. gurneyi (Fuller 1899) [=Eriococcus gurneyi] MG96
         |--A. hakeae (Fuller 1897) [=Eriococcus hakeae] MG96
         |--A. hebes (Hoy 1962) [=Eriococcus hebes] MG96
         |--A. hispidus (Hoy 1962) [=Eriococcus hispidus] MG96
         |--A. humatus (Hoy 1962) [=Eriococcus humatus] MG96
         |--A. iljiniae (Danzig 1972) [=Rhizococcus iljiniae] MG96
         |--A. imperfectus (Fuller 1897) [=Rhizococcus imperfectus] MG96
         |--A. inermis (Green 1915) [=Eriococcus inermis] MG96
         |--A. ironsidei (Williams 1973) [=Eriococcus ironsidei] MG96
         |--A. irregularis (Froggatt 1921) [=Eriococcus irregularis] MG96
         |--A. istriensis (Kozar 1983) [=Gregoporia istriensis] MG96
         |--A. jorgenseni (Morrison 1919) [=Eriococcus jorgenseni] MG96
         |--A. kamahi (Hoy 1958) [=Eriococcus kamahi] MG96
         |--A. kijabensis (James 1934) [=Eriococcus kijabensis] MG96
         |--A. kondarensis (Borchsenius 1949) [=Eriococcus kondarensis] MG96
         |--A. kowhai (Hoy 1962) [=Eriococcus kowhai] MG96
         |--A. laeticoris (Tereznikova 1965) [=Greenisca laeticoris] MG96
         |--A. lanatus (Hempel 1932) [=Eriococcus lanatus] MG96
         |--A. larreae M96
         |--A. latilobatus (Hoy 1962) [=Eriococcus latilobatus] MG96
         |--A. lecanioides (Green 1915) [=Rhizococcus lecanioides] MG96
         |--A. leguminicola (Morrison 1919) [=Eriococcus leguminicola] MG96
         |--A. leptospermi (Maskell 1891) [=Eriococcus leptospermi] MG96
         |--A. lidgetti (Cockerell 1899) [=Rhizococcus lidgetti] MG96
         |--A. mancus (Maskell 1897) [=Rhizococcus casuarinae mancus] MG96
         |--A. maskelli (Hoy 1962) [=Eriococcus maskelli] MG96
         |--A. matai (Hoy 1962) [=Eriococcus matai] MG96
         |--A. matesovae Miller & Gimpel 1996 (see below for synonymy) MG96
         |--A. mendozae (Morrison 1919) [=Eriococcus mendozae] MG96
         |--A. meridianus (Hoy 1962) [=Eriococcus meridianus] MG96
         |--A. milleri (Hoy 1962) [=Eriococcus milleri] MG96
         |--A. mimus (Hoy 1962) [=Eriococcus mimus] MG96
         |--A. montanus (Hoy 1962) [=Eriococcus montanus] MG96
         |--A. montifagi (Hoy 1962) [=Eriococcus montifagi] MG96
         |--A. multispinatus (Tang & Hao 1995) [=Rhizococcus multispinatus] MG96
         |--A. multispinosus (Kuhlgatz 1898) [=Rhizococcus multispinosus] MG96
         |--A. myrsinae (Hoy 1962) [=Eriococcus myrsinae] MG96
         |--A. navarinoensis (Hoy 1962) [=Eriococcus navarinoensis] MG96
         |--A. nelsonensis (Hoy 1962) [=Eriococcus nelsonensis] MG96
         |--A. nematosphaerus (Hu, Xie & Yan 1981) [=Eriococcus nematosphaerus] MG96
         |--A. neomyrti (Hoy 1962) [=Eriococcus neomyrti] MG96
         |--A. ningxianensis (Tang & Hao 1995) [=Eriococcus ningxianensis] MG96
         |--A. nitidulus (Hoy 1962) [=Eriococcus nitidulus] MG96
         |--A. nothofagi (Hoy 1962) [=Eriococcus nothofagi] MG96
         |--A. nuerae (Green 1922) [=Eriococcus nuerae] MG96
         |--A. oblongus (Borchsenius 1949) [=Rhizococcus oblongus] MG96
         |--A. oligacanthus (Danzig 1972) [=Rhizococcus oligacanthus] MG96
         |--A. orariensis (Hoy 1954) [=Eriococcus orariensis] MG96
         |--A. orientalis (Borchsenius 1956) [=Greenisca orientalis] MG96
         |--A. pallidus (Maskell 1885) [=Eriococcus pallidus] MG96
         |--A. palmeri M96
         |--A. papillosus (Morrison 1924) [=Eriococcus papillosus] MG96
         |--A. parabilis (Hoy 1962) [=Eriococcus parabilis] MG96
         |--A. parvisetus [incl. Eriococcus parvispinus Goux 1948] MG96
         |--A. parvulus (Hoy 1962) [=Eriococcus parvulus] MG96
         |--A. paucispinus M96
         |--A. perplexus (Hempel 1900) [=Eriococcus perplexus] MG96
         |--A. philippinensis (Morrison 1920) [=Rhizococcus philippinensis] MG96
         |--A. phyllocladi (Maskell 1892) [=Eriococcus phyllocladi] MG96
         |--A. picta (Froggatt 1916) [=Eriococcus picta] MG96
         |--A. pimeliae (Hoy 1963) [=Eriococcus pimeliae] MG96
         |--A. piptandeniae (Hempel 1937) [=Eriococcus piptandeniae] MG96
         |--A. placidus (Green 1921) [=Eriococcus placidus] MG96
         |--A. podocarpi (Hoy 1962) [=Eriococcus podocarpi] MG96
         |--A. pohutukawa (Hoy 1958) [=Eriococcus pohutukawa] MG96
         |--A. popayanensis (Balachowsky 1959) [=Eriococcus popayanensis] MG96
         |--A. pseudinsignis (Green 1921) [=Eriococcus pseudinsignis] MG96
         |--A. pustulatus (Maskell 1893) [=Rhizococcus pustulatus] MG96
         |--A. puymorensis (Goux 1992) [=Eriococcus puymorensis] MG96
         |--A. quercus M96
         |--A. raithbyi (Maskell 1890) [=Eriococcus raithbyi] MG96
         |--A. rata (Hoy 1958) [=Eriococcus rata] MG96
         |--A. reynei (Schmutterer 1952) [=Eriococcus reynei] MG96
         |--A. rhadinothrix (Miller & González 1975) [=Eriococcus rhadinothrix] MG96
         |--A. rhodomyrti (Green 1922) [=Eriococcus rhodomyrti] MG96
         |--A. rosaceus (Balachowsky 1932) [=Eriococcus rosaceus] MG96
         |--A. rosannae (Tranfaglia & Esposito 1985) [=Eriococcus rosannae] MG96
         |--A. rotundus (Hoy 1962) [=Eriococcus rotundus] MG96
         |--A. rubrus (Matesova 1960) [=Greenisca rubra] MG96
         |--A. rubrifagi (Hoy 1962) [=Eriococcus rubrifagi] MG96
         |--A. rugosus (Froggatt 1933) [=Eriococcus rugosus] MG96
         |--A. rusapiensis (Hall 1937) [=Eriococcus rusapiensis] MG96
         |--A. saboteneus (Kuwana & Tanaka 1922) [=Eriococcus saboteneus] MG96
         |--A. sachalinensis (Siraiwa 1939) [=Eriococcus sachalinensis] MG96
         |--A. salsolae (Borchsenius 1949) [=Rhizococcus salsolae] MG96
         |--A. sanguinairensis (Goux 1993) [=Eriococcus sanguinairensis] MG96
         |--A. serratilobis (Green 1915) [=Eriococcus serratilobis] MG96
         |    |--A. s. serratilobis MG96
         |    `--A. s. prominens (Green 1915) [=Eriococcus serratilobis prominens] MG96
         |--A. setulosus (Hoy 1962) [=Eriococcus setulosus] MG96
         |--A. shiraiwai M96
         |--A. simplex (Maskell 1897) [=Eriococcus simplex] MG96
         |--A. socialis (Goux 1931) [=Eriococcus socialis] MG96
         |--A. sophorae (Green 1929) [=Eriococcus sophorae] MG96
         |--A. sordidus (Green 1904) [=Eriococcus sordidus] MG96
         |--A. spiniger (Maskell 1896) [=Eriococcus spiniger] MG96
         |--A. stenoclini (Mamet 1950) [=Eriococcus stenoclini] MG96
         |--A. sutepensis (Takahashi 19420 [=Eriococcus sutepensis] MG96
         |--A. targassonensis (Goux 1993) [=Eriococcus targassonensis] MG96
         |--A. tavignani (Goux 1991) [=Eriococcus tavignani] MG96
         |--A. tenuis (Green 1922) [=Eriococcus tenuis] MG96
         |--A. tepperi (Maskell 1892) [=Eriococcus tepperi] MG96
         |--A. tesselatus (Froggatt 1916) [=Eriococcus tesselatus] MG96
         |--A. tholothrix (Miller & González 1975) [=Eriococcus tholothrix] MG96
         |--A. thymi (Schrank 1801) [=Coccus thymi] MG96
         |--A. tillandsiae M96
         |--A. timidus (Hulden 1985) [=Eriococcus timidus] MG96
         |--A. tricarinatus (Fuller 1897) [=Eriococcus tricarinatus] MG96
         |--A. tripartitus (Fuller 1897) [=Eriococcus tripartitus] MG96
         |--A. trispinatus (Wang 1974) [=Eriococcus trispinatus] MG96
         |--A. tucurincae (Laing 1929) [=Eriococcus tucurincae] MG96
         |--A. valenzuelae (Balachowsky 1959) [=Eriococcus valenzuelae] MG96
         |--A. variabilis (Goux 1940) [=Eriococcus variabilis] MG96
         |--A. veyrensis (Goux 1991) [=Eriococcus veyrensis] MG96
         |--A. villosus (Froggatt 1916) [=Eriococcus villosus] MG96
         |--A. wangi Miller & Gimpel 1996 [=Eriococcus rugosus Wang 1982 non Froggatt 1933] MG96
         `--A. zygophylli (Archangelskaya 1931) [=Eriococcus zygophylli] MG96

Acanthococcus dziedzickae Miller & Gimpel 1996 [=Rhizococcus palustris Dziedzicka & Koteja 1971 non A. palustris Dodds 1923] MG96

Acanthococcus festucarum (Lindinger 1932) [=Eriococcus festucarum, E. festucae Kuwana & Fukaya in Kuwana 1914 non Targioni Tozzetti 1869] MG96

Acanthococcus matesovae Miller & Gimpel 1996 [=A. multispinosus Matesova 1976 non Rhizococcus multispinosus Kuhlgatz 1898] MG96

*Type species of generic name indicated

REFERENCES

[AY04] Austin, A. D., D. K. Yeates, G. Cassis, M. J. Fletcher, J. La Salle, J. F. Lawrence, P. B. McQuillan, L. A. Mound, D. J. Bickel, P. J. Gullan, D. F. Hales & G. S. Taylor. 2004. Insects 'Down Under'—diversity, endemism and evolution of the Australian insect fauna: examples from select orders. Australian Journal of Entomology 43 (3): 216–234.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[G34] Girault, A. A. 1934. New Capsidae and Hymenoptera, with note on an unmentionable. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 311–314).

[K00] Koteja, J. 2000. Advances in the study of fossil coccids (Hemiptera: Coccinea). Polskie Pismo Entomologiczne 69: 187–218.

[M96] Miller, D. R. 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington 98 (1): 68–86.

[MG96] Miller, D. R., & M. E. Gimpel. 1996. Nomenclatural changes in the Eriococcidae (Homoptera: Coccoidea). Proceedings of the Entomological Society of Washington 98 (3): 597–606.

[MC94] Morrone, J. J., & J. M. Carpenter. 1994. In search of a method for cladistic biogeography: an empirical comparison of component analysis, Brooks parsimony analysis, and three-area statements. Cladistics 10: 99–153.

[N91] Naumann, I. D. 1991. Hymenoptera (wasps, bees, ants, sawflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 916–1000. Melbourne University Press: Carlton (Victoria).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[S07] Ślipiński, A. 2007. Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their biology and classification. Australian Biological Resources Study: Canberra.

[S69] Steyskal, G. C. 1969. The mistreatment of the Latin genitive case in forming names of parasites. Systematic Zoology 18 (3): 339–342.

[W87] Walter, D. E. 1987. Belowground arthropods of semiarid grasslands. In: Capinera, J. L. (ed.) Integrated Pest Management on Rangeland: A shortgrass prairie perspective pp. 271–290. Westview Press, Inc.: Boulder (Colorado).

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

Last updated: 30 April 2022.

Pseudococcidae

Eggplant mealybugs Centrococcus insolitus with ovisacs, from Moore et al. (2014).

Belongs within: Coccoidea.
Contains: Paracoccus, Rhizoecus, Pseudococcus, Nipaecoccus, Phenacoccus.

The Pseudococcidae include the mealy bugs, named for the thin covering of mealy wax over the body surface of many species. Some species are significant plant pests such as the citrus mealybug Planococcus citri. Conversely, the species Hypogeococcus festerianus has been used in Australia to control the harrisia cactus Eriocereus martinii (Carver et al. 1991). Many pseudococcids exhibit highly specialised symbioses with ants, who tend them for the honeydew they produce. Malaicoccus formicarii is found in association with the ant species Hypoclinea cuspidatus, and is transported by its host ants between suitable feeding sites. Emerging queens of the ant genus Acropyga carry mealybugs in their mandibles in order to establish a new mealybug 'herd' when establishing a new ant colony (Grimaldi & Engel 2005). Species of the genus Rhizoecus are subterranean, feeding on the roots of their host plants.

Characters (from Carver et al. 1991): Female with a combination of dorsal ostioles, ventral circuli, trilocular pores and usually 1-18 pairs of cerarii; anterior and posterior pairs of thoracic spiracles approximately the same size; abdominal spiracles absent; terminal segments of abdomen not fused into pygidium; anal ring often with an outer and an inner row of pores; tubular ducts not cupped at inner end.

<==Pseudococcidae
    |--Pseudococcinae [Pseudococcini] A73
    |    |--Pseudococcus GML13
    |    |--Ferrisiana virgata (Cockerell 1893) [=Dactylopius virgatus] A73
    |    `--Nipaecoccus A73
    `--Phenacoccinae [Phenacoccini] KP01
         |--Phenacoccus LS96
         `--Centrococcus insolitus (Green 1908) [=Phenacoccus insolitus] A73

Pseudococcidae incertae sedis:
  Trabutina mannipara GE05
  Hippeococcus GE05
  Malaicococcus formicarii GE05
  Electromyrmococcus GE05
  Tympanococcus Williams 1967 GML13
  Chlorococcus Beardsley 1971 GML13
  Clavicoccus erinaceus Ferris in Zimmerman 1948 GML13
  Phyllococcus oahuensis GML13
  Melanococcus albizziae CGW91
  Australicoccus grevilleae CGW91
  *Ripersiella rumicis M27, WEE70 [=Ripersia rumicis M27]
  Tridiscus distichlii WEE70
  Amonostherium lichtensioides M96
  Anisococcus milleri M96
  Antoninoides parrotti M96
  Ehrhornia cupressi M96
  Geococcus coffeae M96
  Lachnodiella mexicana M96
  Palmicultor palmarum M96
  Prorhizoecus atopoporus M96
  Pseudantonina texana M96
  Radicoccus kelloggi M96
  Saccharicoccus sacchari M96
  Syrmococcus pecosensis M96
  Cryptoripersia arizonensis W87
  Mollicoccus Williams 1960 GML13
    `--M. guadalcanalanus Williams 1960 GML13
  Trionymus M96
    |--T. coronus M96
    `--T. perrisii K09
  Chorizococcus M96
    |--C. rostellum M96
    `--C. triunfoensis M96
  Ferrisia M96
    |--F. consobrina M96
    `--F. virgata M96
  Heliococcus M96
    |--H. corralesi M96
    `--H. deserticola M96
  Porococcus M96
    |--P. pergandei M96
    `--P. tinctorius M96
  Antonina WEE70
    |--A. bambusae B88
    |--A. graminis [incl. A. indica] C81
    `--A. nortoni M96
  Rastrococcus GE05
    |--R. iceryoides GE05
    |--R. invadens MHG04
    `--R. spinosus MHG04
  Humococcus M96
    |--H. atriplicis M96
    |--H. hilariae M96
    `--H. yenseni M96
  Hypogeococcus M96
    |--H. boharti M96
    |--H. festerianus CGW91
    |--H. othnius M96
    `--H. spinosus M96
  Planococcus S07
    |--P. citri A-G91
    |--P. lilacinus MHG04
    |--P. minor CGW91
    `--P. vovae S07
  Distichlicoccus M96
    |--D. megacirculus W87
    |--D. sahuayoensis M96
    |--D. salinus M96
    |--D. sinaloanus M96
    `--D. zacapuensis M96
  Spilococcus M96
    |--S. eriogoni M96
    |--S. larreae M96
    |--S. mamillariae M96
    |--S. sequoiae M96
    `--S. steelii M96
  Cataenococcus M96
    |--C. cualatensis M96
    |--C. ductorum M96
    |--C. guatemalensis M96
    |--C. ingrandi M96
    |--C. larai M96
    `--C. olivaceus M96
  Dysmicoccus MHG04
    |--D. bispinosus M96
    |--D. boninsis M96
    |--D. brevipes MF07
    |--D. grassi M96
    |--D. hurdi M96
    |--D. mackenziei M96
    |--D. neobrevipes MHG04
    |--D. pinicolus M96
    `--D. texensis M96
  Paracoccus M96
  Rhizoecus M96

*Type species of generic name indicated

REFERENCES

[A-G91] Al-Gboory, J. I. 1991. Biology of oriental citrus mite, Eutetranychus orientalis (Klein) on different citrus species. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 2 pp. 607–611. SPB Academic Publishing: The Hague.

[A73] Ali, S. M. 1973. Some coccids from Goa. Journal of the Bombay Natural History Society 69 (3): 669–671.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[C81] Caltagirone, L. E. 1981. Landmark examples in classical biological control. Annual Review of Entomology 26: 213–232.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[GML13] Gullan, P. J., M. L. Moir & M. C. Leng. 2013. A new species of mealybug (Hemiptera: Pseudococcidae) from critically endangered Banksia montana in Western Australia. Records of the Western Australian Museum 28 (1): 13–20.

[K09] Köhler, G. 2009. Heuschrecken (Saltatoria) und Ohrwürmner (Dermaptera) im Immissionsgebiet des Düngemittelwerkes Steudnitz/Thüringen—eine Langzeitstudie (1978–2001). Mauritiana 20 (3): 601–646.

[KP01] Koteja, J., & G. O. Poinar Jr. 2001. A new family, genus, and species of scale insect (Hemiptera: Coccinea: Kukaspididae, new family) from Cretaceous Alaskan amber. Proceedings of the Entomological Society of Washington 103 (2): 356–363.

[LS96] Levins, R., & B. B. Schultz. 1996. Effects of density dependence, feedback and environmental sensitivity on correlations among predators, prey and plant resources: Models and practical implications. Journal of Animal Ecology 65: 802–812.

[MHG04] Maynard, G. V., J. G. Hamilton & J. F. Grimshaw. 2004. Quarantine—phytosanitary, sanitary and incursion management: an Australian entomological perspective. Australian Journal of Entomology 43 (3): 318–328.

[M96] Miller, D. R. 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington 98 (1): 68–86.

[MF07] Moraes, G. J. de, & C. H. W. Flechtmann. 2007. Phytophagous mites of tropical crops in eastern South America. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 279–288. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[M27] Myers, J. G. 1927. On the nomenclature of New Zealand Homoptera. Transactions and Proceedings of the New Zealand Institute 57: 685–690.

[S07] Ślipiński, A. 2007. Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their biology and classification. Australian Biological Resources Study: Canberra.

[W87] Walter, D. E. 1987. Belowground arthropods of semiarid grasslands. In: J. L. Capinera (ed.) Integrated Pest Management on Rangeland: A shortgrass prairie perspective pp. 271–290. Westview Press, Inc.: Boulder (Colorado).

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

Last updated: 30 April 2022.

Pentatomoidea

Aphylum syntheticum, copyright Martin Lagerwey.

Belongs within: Panheteroptera.
Contains: Acanthosomatidae, Scutelleridae, Tessaratomidae, Cydnidae, Pentatominae, Asopinae.

The Pentatomoidea, stink bugs and related taxa, are a group of bugs with an enlarged scutellum, commonly reaching the apex of the clavus or beyond. In extreme cases, the scutellum may cover the entire abdomen. Most pentatomoids are plant-feeders but some species of Pentatomidae are predators. Examples of pentatomoid families in which the scutellum covers the abdomen include the Plataspididae, in which the hemelytra are much longer than the body but folding to fit under the scutellum at rest. Members of the Plataspididae are found in warmer regions of the Old World. Lestonia is an endemic Australian genus that has been collected on growing tips of the native cypress Callitris; members of this genus have the margins of the head, pronotum and part of the costal margins of the hemelytron expanded and recurved to give a tortoise-like appearance (Carver et al. 1991).

Other distinctive members of the Pentatomoidea include the east Asian genus Parastrachia, including two species of large, strikingly red and black bugs with a distinctive subsocial mode of behaviour through progressive provisioning (Sweet & Schaefer 2002). The tropical Urostylinae are relatively elongate and delicate compared to other pentatomoids and characterised by cylindrical antennifers set on the lateral margins of the head, entirely visible from above. The Dinidoridae are an Old World tropical family with reticulate venation on the membrane of the hemelytron (Carver et al. 1991).

<==Pentatomoidea SS02
    |--+--Lestonia China 1955 [Lestoniidae] G75
    |  |    |--*L. haustorifera China 1955 G75
    |  |    `--L. grossi CGW91
    |  `--Plataspididae [Brachyplatidae, Coptosomatidae, Plataspidinae] G75
    |       |--Ceratocoris RD77
    |       |--Elapheozygum RD77
    |       |--Brachyplatys CGW91
    |       |    |--B. pacificus Wall. 1851 D14
    |       |    `--B. vanikorensis Boisduval 1835 B35
    |       `--Coptosoma S07
    |            |--C. ostensum S07
    |            `--C. scutellatum H01
    `--+--Cydnidae G75
       `--+--Phloeidae G75
          `--Pentatomidae G75
               |  i. s.: Erthesina fullo YS01, H01
               |         Irochrotus caspius H01
               |         Phimodera collina H01
               |         Masthletinus abbreviatus H01
               |         Neottiglossa pusilla [incl. N. inflexa] H01
               |         Eusarcoris aeneus H01
               |         Rubiconia intermedia H01
               |         Palomena prasina H01
               |         Picromerus bidens H01
               |         Commius elegans C91
               |         Dalsira bohndorffi RD77
               |         Chlorochroa RD77
               |           |--C. ligata RD77
               |           `--C. uhleri E66
               |         Cyclopelta siccifolia R72
               |         Brochymena E66
               |           |--B. arborea E66
               |           |--B. cariosa E66
               |           |--B. carolinensis E66
               |           `--B. quadripustulata E66
               |         Dendrocoris humeralis E66
               |         Podisus chilensis E66
               |         Stethaulax marmoratus E66
               |         Trichopepla semivittata E66
               |         Solubea pugnax E66
               |--Trincavellius [Discocephalinae] B27
               |--Pentatominae SS02
               |--Aphylum Bergroth 1906 G75 [Aphylidae, Aphylinae CGW91]
               |    |--*A. syntheticum Bergroth 1906 G75
               |    `--A. bergrothi G75
               `--Asopinae SS02

Pentatomoidea incertae sedis:
  Acanthosomatidae SS02
  Parastrachia Distant 1883 SDT88 [Parastrachiaria, Parastrachiidae SS02, Parastrachiinae]
    |--P. japonensis (Scott 1880) (see below for synonymy) SDT88
    `--P. nagaensis Distant 1908 SS02
  Dinidoridae SS02
    |--Coridius janus SS02
    |--Eumenotes obscurus CGW91
    `--Megymenum SS02
         |--M. affine Boisduval 1835 B35
         |--M. dentatum Boisduval 1835 B35
         |--M. gracilicorne SS02
         `--M. insulare CGW91
  Thyreocoridae SS02
    |--Thyreocoris SS02
    `--Corimelaena [Corimelaeninae] SS02
  Scutelleridae GE05
  Tessaratomidae SS02
  Megaris [Megarididae] SDT88
  Canopidae CGW91
  Urostylinae [Urolabidae] SDT88
    |--Urolabida nilgirica SDT88
    |--Urochela pardalina Zhang 1989 RJ93
    `--Urostylis RD77
  Serbana SDT88
  Cyrtocoridae SDT88
  Ruckesona [Saileriolinae] SDT88
    `--R. vitrella SDT88
  Tahitocoris SDT88

Parastrachia japonensis (Scott 1880) [=Asopus japonensis, P. japonica (l. c.); incl. *P. fulgens Distant 1883] SDT88

*Type species of generic name indicated

REFERENCES

[B27] Bergroth, E. 1927. Hemiptera Heteroptera from New Zealand. Transactions and Proceedings of the New Zealand Institute 57: 671–684.

[B35] Boisduval, J. B. 1835. Voyage de Découvertes de l’Astrolabe. Exécuté par ordre du Roi, pendant les années 1826–1827–1828–1829, sous le commandement de M. J. Dumont d'Urville. Faune entomologique de l'océan Pacifique, avec l'illustration des insectes nouveaux recueillis pendant le voyage vol. 2. Coléoptères et autres ordres. J. Tastu: Paris.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[C91] CSIRO. 1991. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 1. Melbourne University Press: Carlton (Victoria).

[D14] Distant, W. L. 1914. Rhynchota from New Caledonia and the surrounding islands. In: Sarasin, F., & J. Roux (eds) Nova Caledonia: Forschungen in Neu-Caledonian und auf den Loyalty-Inseln. A. Zoologie vol. 1 pt 4 pp. 369–390, pls 11–12. C. W. Kreidels Verlag: Wiesbaden.

[E66] Evans, H. E. 1966. The Comparative Ethology and Evolution of the Sand Wasps. Harvard University Press: Cambridge (Massachusetts).

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G75] Gross, G. F. 1975. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera—Part I. Handbook of the Flora and Fauna of South Australia.

[H01] Horváth, G. 1901. Hemipterák [Hemiptera]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 245–274. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[R72] Rao, K. R. 1972. On the occurrence of Cyclopelta siccifolia Westw. (Hemiptera: Pentatomidae) on Zizyphus sp. in Aurangabad. Journal of the Bombay Natural History Society 69 (2): 440–441.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[SDT88] Schaefer, C. W., W. R. Dolling & S. Tachikawa. 1988. The shieldbug genus Parastrachia and its position within the Pentatomoidea (Insecta: Hemiptera). Zoological Journal of the Linnean Society 93: 283–311.

[S07] Ślipiński, A. 2007. Australian Ladybird Beetles (Coleoptera: Coccinellidae): Their biology and classification. Australian Biological Resources Study: Canberra.

[SS02] Sweet, M. H., & C. W. Schaefer. 2002. Parastrachiinae (Hemiptera: Cydnidae) raised to family level. Annals of the Entomological Society of America 95: 441–448.

[YS01] Yoshizawa, K., & T. Saigusa. 2001. Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera) based on forewing base structure with comments on monophyly of Auchenorrhycha (Hemiptera). Systematic Entomology 26: 1–13.

Last updated: 9 May 2022.

Coccidae

Fig wax scale Ceroplastes rusci, from here.

Belongs within: Coccoidea.
Contains: Lecanium, Coccus.

The Coccidae, soft scales, include both wax-covered and naked species, characterised by the presence in females of paired triangular or rounded plates at the base of an anal cleft. Several species are significant crop pests such as the hemispherical scale Saissetia coffeae and the soft wax scale Ceroplastes destructor.

Characters (from Carver et al. 1991): Female with paired triangular or rounded anal plates at the base of an anal cleft; anterior and posterior thoracic spiracles approximately the same size; spiracular and marginal setae present; tubular ducts with inner end cupped.

<==Coccidae [Lecaniidae]
    |--Lecaniinae H01b
    |    |--Lecanium G25
    |    `--Pseudokermes H01b
    |         |--P. armatus M96
    |         `--P. nitens H01b
    |--Coccinae [Coccini] A73
    |    |--Coccus L58
    |    `--Saissetia A73
    |         |--S. coffeae CGW91 [=Lecanium coffeae B88]
    |         |--S. hemisphaerica (Targioni 1867) [=Lecanium hemisphaericum] A73
    |         |--S. miranda M96
    |         |--S. neglecta M96
    |         |--S. nigra B88
    |         |--S. oleae B88
    |         `--S. tolucana M96
    `--Ceroplastes [Ceroplastinae] A73
         |--C. actiniformis Green 1896 A73
         |--C. africanus [incl. C. africanus var. cristatus Green 1899, C. egbarum Cockerell 1899] G00
         |--C. albolineatus M96
         |    |--C. a. albolineatus M96
         |    `--C. a. volcanicus M96
         |--C. amazonicus Hempel 1901 H01c
         |--C. angulatus M96
         |--C. brachyurus M96
         |--C. ceriferus B88
         |--C. chrysanthemum G17
         |--C. cirripediformis M96
         |--C. cistudiformis M96
         |--C. coloratus M96
         |--C. communis Hempel 1901 H01c
         |--C. destructor CGW91
         |--C. dugesii M96
         |--C. floridensis M96
         |--C. grandis Hempel 1901 H01c
         |--C. irregularis M96
         |--C. macgregori M96
         |--C. marmoreus M96
         |--C. minutus M96
         |--C. novaesi Hempel 1901 H01c
         |--C. purpurellus M96
         |--C. rubens CGW91
         |--C. rusci B88
         |--C. sinensis M96
         `--C. variegatus Hempel 1901 H01c

Coccidae incertae sedis:
  Parasaissetia nigra CGW91
  Phenacaspis dubia [=Chionaspis dubia] M27
  Eulecanium GE05
    |--E. coryli RD77
    |--E. patersoniae GE05
    `--E. subaustrale M96
  Symonicoccus CGW91
  Cryptes baccatus CGW91
  Pulvinaria M96
    |--P. elongata CGW91
    `--P. peninsularis M96
  Antecerococcus bryoides K08
  Brachyscelis duplex P08
  Paralecanium expansum WEE70
  Ericerus pela WEE70
  Platycoelostoma Morrison 1923 M27
    `--*P. compressa [=Coelostomidia compressa] M27
  Anoplaspis M27
    |--*A. metrosideri [=Lepidosaphes metrosideri] M27
    `--A. maskelli Morrison 1922 M27
  Ctenochiton B88
    |--C. aztecus M96
    |--C. elaeocarpi B88
    |--C. perforatus B88
    `--C. viridis B88
  Solenococcus H01a
    |--S. baccharidis Hempel 1901 H01a
    `--S. tuberculus Hempel 1901 H01a
  Stigmacoccus Hempel 1901 H01a
    `--*S. asper Hempel 1901 H01a
  Apiococcus Hempel 1901 H01a
    |--*A. gregarius Hempel 1901 H01a
    |--A. asperatus Hempel 1901 H01a
    |--A. globosus Hempel 1901 H01a
    `--A. singularis Hempel 1901 H01a
  Tectococcus Hempel 1901 H01a
    `--*T. ovatus Hempel 1901 H01a
  Akermes M96
    |--A. colimae M96
    `--A. townsendi M96
  Cardiococcus umbonatum M96
  Ceroplastodes dugesii M96
  Inglisia malvacearum M96
  Kilifia M96
    |--K. acuminata M96
    `--K. americana M96
  Megasaissetia inflata M96
  Mesolecanium M96
    |--M. impar M96
    |--M. nocturnum M96
    `--M. perditum M96
  Metapulvinaria lycii M96
  Milviscutulus mangiferae M96
  Neolecanium M96
    |--N. chilaspidis M96
    |--N. herrerae M96
    |--N. imbricatum M96
    |--N. manzanillense M96
    |--N. plebeium M96
    |--N. sallei M96
    `--N. tuberculatum M96
  Pendularia jaliscensis M96
  Philephedra M96
    |--P. colimensis M96
    |--P. crescentiae M96
    |--P. lutea M96
    |--P. mimosae M96
    |--P. parvula M96
    `--P. tuberculosa M96
  Platysaissetia castilloae M96
  Protopulvinaria pyriformis M96
  Schizochlamidia mexicana M96
  Tillanococcus M96
    |--T. mexicana M96
    `--T. tillandsiae M96
  Toumeyella M96
    |--T. cerifera M96
    |--T. mirabilis M96
    `--T. sonorensis M96
  Insulaspis pallidula CK15

*Type species of generic name indicated

REFERENCES

[A73] Ali, S. M. 1973. Some coccids from Goa. Journal of the Bombay Natural History Society 69 (3): 669–671.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[CK15] Calvo, F. J., M. Knapp, Y. M. van Houten, H. Hoogerbrugge & J. E. Belda. 2015. Amblyseius swirskii: what made this predatory mite such a successful biocontrol agent? Experimental and Applied Acarology 65: 419–433.

[CGW91] Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera (bugs, leafhoppers, cicadas, aphids, scale insects etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 429–509. Melbourne University Press: Carlton (Victoria).

[G17] Girault, A. A. 1917. Descriptiones hymenopterorum chalcidoidicarum variorum cum observationibus. V. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 116–131).

[G25] Girault, A. A. 1925. An essay on when a fly is lovable, the ceremony of baptizing some and unlovely hate. Privately published (reprinted: Gordh, G., A. S. Menke, E. C. Dahms & J. C. Hall. 1979. The privately printed papers of A. A. Girault. Memoirs of the American Entomological Institute 28: 187–190).

[G00] Green, E. E. 1900. Note on Ceroplastes africanus (family Coccidae). Annals and Magazine of Natural History, series 7, 5: 158–159.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[H01a] Hempel, A. 1901a. Descriptions of Brazilian Coccidae. Annals and Magazine of Natural History, series 7, 7: 110–125.

[H01b] Hempel, A. 1901b. Descriptions of Brazilian Coccidae (continued). Annals and Magazine of Natural History, series 7, 7: 206–219.

[H01c] Hempel, A. 1901c. Descriptions of Brazilian Coccidae (continued). Annals and Magazine of Natural History, series 7, 7: 556–561.

[K08] Kirkaldy, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proceedings of the Linnean Society of New South Wales 33: 345–391, pl. 4.

[L58] Linnaeus, C. 1758. Systema Naturae per Regna Tria Naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii: Holmiae.

[M96] Miller, D. R. 1996. Checklist of the scale insects (Coccoidea: Homoptera) of Mexico. Proceedings of the Entomological Society of Washington 98 (1): 68–86.

[M27] Myers, J. G. 1927. On the nomenclature of New Zealand Homoptera. Transactions and Proceedings of the New Zealand Institute 57: 685–690.

[P08] Palmer. 1908. Notes and exhibits. Proceedings of the Linnean Society of New South Wales 33: 434.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[WEE70] Woodward, T. E., J. W. Evans & V. F. Eastop. 1970. Hemiptera (bugs, leafhoppers, etc.) In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 387–457. Melbourne University Press.

Last updated: 30 April 2022.