Blattoidea

Cretaceous predatory roach Raphidiomimula burmitica, from Grimaldi & Ross (2004).

Belongs within: Dictyoptera.
Contains: Mantodea, Isoptera, Blattaria.

The cockroach superfamily Blattoidea, as used by Vršanský et al. (2002), is paraphyletic to Polyphagoidea, Mantodea and Isoptera, and hence includes all crown-group dictyopterans and some related fossil families. These include the Raphidiomimidae, a Late Jurassic to Cretaceous group of predatory roaches with long slender legs, of which the front pair are raptorial (Grimaldi & Ross 2004). They also include the Upper Jurassic Skok svaba, which exhibited adaptations for jumping (Bohn et al. 2010).

Synapomorphies (from Vršanský et al. 2002): Ovipositor very short externally; eggs laid in two-row packages (in crown taxa at least); size small (body length under 20 mm, fore wing about 8-15 mm); fore wing elongate, with fore and hind margins parallel.

<==Blattoidea [Polyphagoidea] VVR02
    |--+--Blattulidae GE05
    |  |    |--Elisama vidlickai Vršanský 2004 V10
    |  |    `--Tarakanula shcherbakovi VLR09
    |  `--+--+--Liberiblattina GR04 [Liberiblattinidae V10]
    |     |  |    `--L. ihringovae GR04
    |     |  |--+--Eadiidae V10
    |     |  |  `--Mantodea GE05
    |     |  |--Skok [Skokidae] V10
    |     |  |    `--S. svaba Vršanský 2007 BP10
    |     |  `--Isoptera V10
    |     `--Blattaria GE05
    `--+--Mesoblattinidae GE05
       |    |--Mesoblattina protypa VVR02
       |    |--Hispanoblatta sumptuosa VVR02
       |    |--Artitocoblatta asiatica VVR02
       |    `--Austroblattula F71
       `--Raphidiomimidae GE05
            |--Raphidiomima Vishniakova 1973 GR04
            |    |--R. chimaera GR04
            |    `--R. cognata VVR02
            |--Cretaceochorista Z02
            |--Rhipidoblattina katavica GE05
            |--Raphidiomimula Grimaldi & Ross 2004 GR04
            |    `--*R. burmitica Grimaldi & Ross 2004 GR04
            `--Cameloblatta variegata GR04

*Type species of generic name indicated

REFERENCES

[BP10] Bohn, H., M. Picker, K.-D. Klass & J. Colville. 2010. A jumping cockroach from South Africa, Saltoblattella montistabularis, gen. nov., spec. nov. (Blattodea: Blattellidae). Arthropod Systematics and Phylogeny 68 (1): 53-69.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

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

[GR04] Grimaldi, D. A. & A. J. Ross. 2004. Raphidiomimula, an enigmatic new cockroach in Cretaceous amber from Myanmar (Burma) (Insecta: Blattodea: Raphidiomimidae). Journal of Systematic Palaeontology 2: 101-104.

[V10] Vršanský, P. 2010. Cockroach as the earliest eusocial animal. Acta Geologica Sinica (English Edition) 84 (4): 793-808.

[VLR09] Vršanský, P., J.-H. Liang & D. Ren. 2009. Advanced morphology and behaviour of extinct earwig-like cockroaches (Blattida: Fuziidae fam. nov.). Geologica Carpathica 60 (6): 449-462.

[VVR02] Vršanský, P., V. N. Vishniakova & A. P. Rasnitsyn. 2002. Order Blattida Latreille, 1810. The cockroaches (=Blattodea Brunner von Wattenvill, 1882). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 263-270. Kluwer Academic Publishers: Dordrecht.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

Insecta

Fossil of Dasyleptus sp. from the Mazon Creek, from RCFossils.

Belongs within: Panhexapoda.
Contains: Zygentoma, Pterygota, Meinertellidae, Machilidae.

One of the most speciose of all groups of organisms, the insects dominate the terrestrial environment. The majority of insects belong to the winged clade Pterygota, but some basal clades retain the primitive wingless morphology. These latter groups, the Archaeognatha and Zygentoma, are superficially similar in having a more or less elongate body form with three long terminal filaments (the cerci and median filament), and so have been combined in the past under the name Thysanura. However, the Zygentoma share a number of synapomorphies with the Pterygota, including the presence of mandibles with a secondary articulation in addition to the single articulation found in Archaeognatha, and are now recognised a forming a clade with the winged insects (Grimaldi & Engel 2005). The Archaeognatha, bristletails, are characterised by large eyes that meet on the top of the head and a distinctly humped thorax. They jump by rapidly flexing the abdomen to propel themselves into the aire (Grimaldi & Engel 2005).

Synapomorphies (from Grimaldi & Engel 2005, many altered in derived subgroups): Antenna lacking musculature beyond the scape; chordotonal organ (Johnston's organ) present in antennal pedicel; posterior tentorium developed into transverse bar; articulations absent between coxae and sterna; tarsus subsegmented into tarsomeres; pretarsal claws articulating with apicalmost tarsomere rather than with tarsal base; ovipositor present in females formed from gonapophyses on eighth and ninth abdominal segments; long terminal filament present on dorsum of eleventh abdominal segment.

<==Insecta [Ectognatha, Ectotropha, Lepismatona, Scarabaeoda, Thysanura, Thysanuroidea]
    |--Dicondylia [Cercofilata] GE05
    |    |--Tricholepidion gertschi Wygodzinsky 1961 TW05, KZ02
    |    `--+--Zygentoma TW05
    |       `--Pterygota TW05
    `--Archaeognatha [Machilida, Machiloidea, Microcoryphia, Monocondylia] TW05
         |  i. s.: Triassomachilis Sharov 1948 KS02 [Triassomachilidae GE05]
         |           `--T. uralensis GE05
         |--Dasyleptus Brongniart 1885 FT05 [Cercopodata, Dasyleptidae R02, Monura]
         |    |--D. brongniarti Sharov 1957 K-P83
         |    `--D. sharovi GE05
         `--+--Meinertellidae R02
            `--Machilidae R02

Insecta incertae sedis:
  Drosicha mangiferae G01
  Clavigralla gibbosa G01
  Ramsdelepidion Kukalová-Peck 1987 K-P87
    `--*R. schusteri Kukalová-Peck 1987 K-P87
  Gaspea palaeoentognathae Z02
  Biorhiza pallida A71
  Neocolochelyna itoi Takeuchi 1951 I92
  Paratlanticus tsushimensis Yamasaki 1986 I92
  Cercomantispa shirozui Nakahara 1961 I92
  Armatocillenus sumaoi (Morita 1981) I92
  Heplochlaenius insularis Uéno 1964 I92
  Hydrochus chubu Balfour-Browne & Satô 1962 I92
  Georissus granulosus Satô 1972 I92
  Liparocephalus tokunagai Sakaguchi 1944 I92
  Schwarzerium viridicyaneum Hayashi 1956 I92
  Baryrhynchus tokarensis Ohbayashi & Satô 1966 I92
  Cyphagogus iwatensis Morimoto 1976 I92
  Ogasawarazo daitoensis Voss 1971 I92
  Xyelecia japonica Tagashi 1972 I92
  Pleroneura hikosana Togashi 1972 I92
  Nipponorhynchus I92
    |--N. bimaculatus Naito 1973 I92
    `--N. mirabilis Takeuchi 1941 I92
  Caenosclerogibba japonica Yasumatsu 1958 I92
  Pararrhynchium tsunekii Tano & Yamane 1983 I92
  Liacos melanogaster Tsuneki 1982 I92
  Strymonidia iyonis Ota & Kusunoki 1957 I92
  Pithecops I92
    |--P. corvus I92
    |    |--P. c. corvus I92
    |    `--P. c. ryukyuensis Shirôzu 1964 I92
    `--P. fulgens I92
         |--P. f. fulgens I92
         `--P. f. tsushimanus Shirôzu & Urata 1957 I92
  Tongeia fischeri (Eversmann 1843) I92
  Archiearis notha (Hübner 1808) I92
  Heliophobus texturatus (Alpheraky 1892) I92
  Cratosomus flavofasciatus H91
  Eldana saccharina H91
  Maruca testulalis H91
  Tricopria drosophilae GJR97
  Asobara tabida GJR97
  Aramigus tesselatus GJR97
  Leptopilina GJR97
    |--L. australis GJR97
    `--L. clavipes GJR97
  Schizaphis graminum SF98
  Leptophyllus Hope 1842 FT93
  Amaurocoris laticeps G89
  Aspongopus viduatus G89
  Nariscus spinosus G89
  Spilostethus G89
    |--S. macilentus G89
    `--S. militaris G89
  Anepsius encaustus G89
  Ischnopeza hirticornis G89
  Piezostethus galactinus G89
  Samea multiplicalis B04
  Neohydronomus affinus MHG04
  Araecerus fasciculatus G17a
  Lecanium G25
    |--L. baccatum G25
    |--L. capraea G17d
    `--L. viride G17a
  Stictococcus gowdeyi G17b
  Tachardia decorella G17b
  Physokermes insignicola G17b
  Eutettix tenella G17b
  Dibolia borealis G17b
  Tephrocerus gracilis G17b
  Evetria taxifoliella G17c
  Dioryctria xanthaenobares G17c
  Chelymorpha argus G17c
  Eudamus proteus G18
  Melitara junctolineella B88
  Moneilema ulkei G25
  Mytilaspis cordylinidis G29
  Cylindrococcus B88
    |--C. amplior G34
    `--C. casuarinae B88
  Proceras sacchariphagum B88
  Hypolycaena phorbas B88
  Stomopteryx simplexella B88
  Diplosis sorghicola B88
  Tryporyza VHK02
    |--T. incertulas B88
    |--T. monostigma VHK02
    |--T. nivella VHK02
    `--T. rhodoproctalis VHK02
  Actinoscytina belmontensis Tillyard 1926 F71
  Adelidium cordatum Tillyard 1917 F71
  Agetochoristella adscita Riek 1952 F71
  Anomaloscytina F71
    |--A. incompleta Evans 1943 F71
    `--A. metapteryx Davis 1942 F71
  Aphryganoneura anomala Tillyard 1926 F71
  Archaeosmylus pectinatus Riek 1952 F71
  Atelophlebis culleni (Etheridge & Olliff 1890) [=Ephemera culleni] F71
  Austroidelia perplexa Riek 1954 F71
  Austroprosbole maculata Evans 1943 F71
  Austroscytina imperfecta Evans 1943 F71
  Beaconiella F71
    |--B. fennahi Evans 1963 F71
    `--B. multivenata Evans 1963 F71
  Belpsylla reticulata Evans 1943 F71
  Clavopsyllidium minutum Davis 1942 F71
  Elateridium angustius Tillyard 1917 F71
  Eochiliocycla angusta Davis 1942 F71
  Eopsyllidium delicatulum Davis 1942 F71
  Eupincombea postiva Davis 1942 F71
  Homaloscytina plana Tillyard 1926 F71
  Mesonotoperla sinuata Riek 1954 F71
  Mesophlebia antinodalis Tillyard 1916 F71
  Mesothyris westraliensis Riek 1968 F71
  Metrorhynchites grandis Tillyard 1917 F71
  Mitchelloneura permiana Tillyard 1921 F71
  Nannochoristella reducta Riek 1953 F71
  Neoageta elongata Riek 1953 F71
  Neochoristella aptata Riek 1953 F71
  Neopetromantis australis Riek 1953 F71
  Notoblattites F71
    |--N. mitchelli Tillyard 1917 F71
    `--N. wianamattensis Tillyard 1917 F71
  Orthoscytina F71
    |--O. belmontensis Tillyard 1926 F71
    |--O. indistincta Tillyard 1926 F71
    |--O. mitchelli Tillyard 1926 F71
    |--O. obliqua Tillyard 1926 F71
    |--O. pincombei Tillyard 1926 F71
    |--O. quinquemedia Tillyard 1926 F71
    |--O. subcostalis Tillyard 1926 F71
    `--O. tetraneura Tillyard 1926 F71
  Palaeovicia incerta Evans 1943 F71
  Permagra distincta Evans 1943 F71
  Permopsychops belmontensis Tillyard 1926 [incl. Permithone venosa Davis 1943] F71
  Permoberothella perplexa Riek 1953 F71
  Permobrachus F71
    |--P. dubia (Tillyard 1926) [=Permodiphthera dubia] F71
    `--P. magnus Evans 1943 F71
  Permocapitus globulus Evans 1943 F71
  Permocephalus knighti Evans 1943 F71
  Permodiphthera robusta Tillyard 1926 F71
  Permoglyphis belmontensis Tillyard 1926 F71
  Permojassus F71
    |--P. australis Tillyard 1926 F71
    `--P. dubius Tillyard 1926 F71
  Permopsyllidium F71
    |--P. affine Tillyard 1926 F71
    `--P. mitchelli Tillyard 1926 F71
  Permopsyllidops stanleyi Davis 1942 F71
  Permopsylloides insolita Evans 1943 F71
  Permorapisma biserialis Tillyard 1926 [incl. P. triserialis Tillyard 1926] F71
  Permoscarta F71
    |--P. mitchelli Tillyard 1917 F71
    `--P. triventulata Tillyard 1926 F71
  Permosyne F71
    |--P. affinis Tillyard 1924 F71
    |--P. belmontensis Tillyard 1924 F71
    |--P. mitchelli Tillyard 1924 F71
    `--P. pincombeae Tillyard 1924 F71
  Permothea latipennis Tillyard 1926 F71
  Permotheela scytinopteroides Davis 1942 F71
  Permovicia obscura Evans 1943 F71
  Phipoides elegans Riek 1953 F71
  Prohagla superba Riek 1954 F71
  Protopincombea obgscura Evans 1943 F71
  Protopsyllops minuta Evans 1943 F71
  Psocopsyllidium media Davis 1942 F71
  Psocoscytina bifida Davis 1942 F71
  Psyllidella magna Evans 1943 F71
  Psyllidiana davisia Evans 1943 F71
  Stanleyana pulchra Evans 1943 F71
  Stenoglyphis kimblensis Evans 1947 F71
  Stenoscytina australiensis Tillyard 1926 F71
  Triassagrion australiensis Tillyard 1922 F71
  Triassodoecus chinai Evans 1963 F71
  Triassophlebis stigmata Tillyard 1922 F71
  Triassopsylla plecioides Tillyard 1917 F71
  Triassocytinopsis paranotalis Evans 1956 F71
  Tripsyllidium wadei Evans 1956 F71
  Tychticoloides belmontensis Evans 1963 F71
  Tenebroides mauritanicus BC91
  Fumea casta BC91
  Paleacrita vernata Z91
  Itame argillacearia DC10
  Biloba subsecivela VHK02
  Diaphania indica [=Margaronia indica] VHK02
  Lygropia quaternalis VHK02
  Marasma suspicalis VHK02
  Psara stultalis VHK02
  Holcocera pulverea VHK02
  Noorda moringa VHK02
  Nephanteryx rhodobasalis VHK02
  Anomalococcus indicus VHK02
  Phidodonta modesta VHK02
  Bissetia steniella VHK02
  ‘Cheyletus’ nigripes Mola 1907 B49
  Stephanostoma Danielsen 1880 D56
  Phrynomorphus Curtis 1831 FG06

*Type species of generic name indicated

REFERENCES

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[B49] Baker, E. W. 1949. A review of the mites of the family Cheyletidae in the United States National Museum. Proceedings of the United States National Museum 99 (3238): 267-320.

[BC91] Baker, G. T., & A. Chandrapatya. 1991. Fine structure of the chorion of the broak mite Polyphagotarsonemus latus (Prostigmata: Tarsonemidae). 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. 325-328. SPB Academic Publishing: The Hague.

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

[B04] Briese, D. T. 2004. Weed biological control: applying science to solve seemingly intractable problems. Australian Journal of Entomology 43 (3): 304-317.

[D56] Dawes, B. 1956. The Trematoda with special reference to British and other European forms. University Press: Cambridge.

[DC10] Drummond, F. A., J. A. Collins, B. Choate, D. Woodman, D. T. Jennings, H. Y. Forsythe, Jr & J. C. Cokendolpher. 2010. Harvestman (Opiliones) fauna associated with Maine lowbush blueberry fields in the major production areas of Washington and Hancock counties. Environmental Entomology 39 (5): 1428-1440.

[FT05] Fayers, S. R., & N. H. Trewin. 2005. A hexapod from the Early Devonian Windyfield Chert, Rhynie, Scotland. Palaeontology 48 (5): 1117-1130.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i-viii, 1-351.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[FG06] Frost, D. R., T. Grant, J. Faivovich, R. H. Bain, A. Haas, C. F. B. Haddad, R. O. de Sá, A. Channing, M. Wilkinson, S. C. Donnellan, C. J. Raxworthy, J. A. Campbell, B. L. Blotto, P. Moler, R. C. Drewes, R. A. Nussbaum, J. D. Lynch, D. M. Green & W. C. Wheeler. 2006. The amphibian tree of life. Bulletin of the American Museum of Natural History 297: 1-370.

[G89] Gestro, R. 1889. Viaggio ab Assab nel Mar Rosso dei signori G. Doria ed O. Beccari con il R. Avviso «Esploratore» dal 16 Novembre 1879 al 26 Febbraio 1880.—IV. Coleotteri. Annali del Museo Civico di Storia Naturale di Genova, Serie 2^a 7: 5-72.

[GJR97] Giordano, R., J. J. Jackson & H. M. Robertson. 1997. The role of Wolbachia bacteria in reproductive incompatibilities and hybrid zones of Diabrotica beetles and Gryllus crickets. Proceedings of the National Academy of Sciences of the USA 94: 11439-11444.

[G17a] Girault, A. A. 1917a. New Javanese Hymenoptera. 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: 59-71).

[G17b] Girault, A. A. 1917b. 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).

[G17c] Girault, A. A. 1917c. Descriptiones hymenopterorum chalcidoidicarum variorum cum observationibus. III. 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: 102-111).

[G17d] Girault, A. A. 1917d. 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).

[G18] Girault, A. A. 1918. North American Hymenoptera Trichogrammatidae. 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: 142-152).

[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).

[G29] Girault, A. A. 1929. Description of a case of lunacy in Homo and of six-legged articulates. 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: 262-265).

[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).

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

[G01] Gupta, S. K. 2001. A conspectus of natural enemies of phytophagous mites and mites as potential biocontrol agents of agricultural pests in India. In Acarology: Proceedings of the 10^th International Congress (R. B. Halliday, D. E. Walter, H. C. Proctor, R. A. Norton & M. J. Colloff, eds) pp. 484-497. CSIRO Publishing: Melbourne.

[H91] Hokkanen, H. M. T. 1991. Trap cropping in pest management. Annual Review of Entomology 36: 119-138.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KZ02] Klass, K.-D., O. Zompro, N. P. Kristensen & J. Adis. 2002. Mantophasmatodea: A new insect order with extant members in the Afrotropics. Science 296: 1456-1459.

[KS02] Kluge, N. Yu., & N. D. Sinitshenkova. 2002. Order Ephemerida Latreille, 1810. The true mayflies (=Ephemeroptera Hyatt et Arms, 1891 (s. l.); =Euephemeroptera Kluge, 2000. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 89-97. Kluwer Academic Publishers: Dordrecht.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618-1669.

[K-P87] Kukalová-Peck, J. 1987. New Carboniferous Diplura, Monura, and Thysanura, the hexapod ground plan, and the role of thoracic side lobes in the origin of wings (Insecta). Canadian Journal of Zoology 65: 2327-2345.

[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.

[R02] Rasnitsyn, A. P. 2002. Subclass Lepismatona Latreille, 1804. The wingless insects (=Thysanura Latreille 1796, s. l.) In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 69-74. Kluwer Academic Publishers: Dordrecht.

[SF98] Saikkonen, K., S. H. Faeth, M. Helander & T. J. Sullivan. 1998. Fungal endophytes: a continuum of interactions with host plants. Annual Review of Ecology and Systematics 29: 319-343.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240-257.

[VHK02] Verma, M., M. Hayat & S. I. Kazmi. 2002. The species of Elasmus from India (Hymenoptera: Chalcidoidea: Eulophidae). Oriental Insects 36: 245-306.

[Z91] Zhang, Z. Q. 1991. Biology of mites of Allothrombiinae (Acari: Trombidiidae) and their potential role in pest control. 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. 513-520. SPB Academic Publishing: The Hague.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

Coccoidea

Female and male of Apiomorpha, copyright Galler18.

Belongs within: Coccidomorpha.
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).

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
    |--Sucinikermes Koteja 1988 K00
    |    `--S. kulickae Koteja 1988 K00
    |--Allokermes M96
    |    |--A. galliformis M96
    |    `--A. grandis M96
    `--Olliffiella secunda 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 H91
    `--D. tomentosus M96
  Apiomorpha B88 [Apiomorphidae K00]
    |--A. conica G84
    |--A. duplex AY04
    |--A. frenchi B88
    |--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. 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
  Stictococcus G17 [Stictococcidae CGW91]
    `--S. gowdeyi G17
  Asterolecaniidae CGW91
  Kerriidae [Lacciferidae] CGW91
    |--Kerria lacca GE05
    |--Austrotachardia angulata CGW91
    |--Tachardina CGW91
    `--Laccifer lacca WEE70
  Conchaspis [Conchaspididae] CGW91
    |--C. angraeci CGW91
    `--C. newsteadi M96
  Tachardiidae M96

*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).

[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.

[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.

[H91] Howarth, F. G. 1991. Environmental impacts of classical biological control. Annual Review of Entomology 36: 485–509.

[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).

[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: 21 December 2019.

Coccinea

Cottony cushion scale Icerya purchasi, from Clemson University.

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

The Coccinea include the scale insects, a characteristic 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.

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
    |         Gascardia destructor DAF07
    |         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] B88
                              |    |--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).

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

[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: 8 February 2020.

Eriococcidae

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

Belongs within: Coccoidea.

The Eriococcidae include scale insects causing sometimes remarkable galls on trees.

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
    |--Madarococcus MC94
    |--Opisthoscelis AY04
    |--Cystococcus AY04
    |--Lachnodius eucalypti B88
    |--Keithia Koteja 2000 K00
    |    `--K. luzzii Koteja 2000 K00
    |--Gedanicoccus Koteja 1988 K00
    |    `--G. gracilis Koteja 1988 K00
    |--Kuenowicoccus Koteja 1988 K00
    |    `--K. pietrzeniukae Koteja 1988 K00
    |--Eriococcus B88
    |    |--E. coriaceus B88
    |    `--E. nitidus B88
    |--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

*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).

Carver, M., G. F. Gross & T. E. Woodward. 1991. Hemiptera. In: CSIRO. The Insects of Australia, 2nd ed., vol. 1, pp. 429-509. Melbourne University Press.

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

[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.

Pseudococcidae

Long-tailed mealy bug Pseudococcus longispinus, photographed by James Castner.

Belongs within: Coccoidea.

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 Pseudococcus longispinus on citrus and the polyphagous P. affinis (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
    |--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
    |--Nipaecoccus M96
    |    |--N. aurilanatus B88
    |    |--N. cercidii M96
    |    |--N. filamentosus M96
    |    |--N. filicis M96
    |    |--N. gilli M96
    |    |--N. jonmartini M96
    |    |--N. kosztaraborum M96
    |    |--N. neogaeus M96
    |    |--N. nipae M96
    |    |--N. vastator A-G91
    |    `--N. zapotlanus M96
    |--Paracoccus M96
    |    |--P. alazanensis M96
    |    |--P. baccharidicola M96
    |    |--P. circuliprivis M96
    |    |--P. ferrisi M96
    |    |--P. hamoni M96
    |    |--P. herreni M96
    |    |--P. juniperi M96
    |    |--P. lycopersici M96
    |    |--P. marginatus M96
    |    |--P. mexicanus M96
    |    |--P. oneratus M96
    |    |--P. ordinis M96
    |    |--P. reductus M96
    |    |--P. salviacola M96
    |    |--P. solani M96
    |    |--P. townsendi M96
    |    `--P. villanuevai M96
    |--Pseudococcus Westwood 1840 GML13
    |    |--*P. longispinus (Targioni Tozzetti 1867) [=Dactylopius longispinus] GML13
    |    |--P. adonidum B88
    |    |--P. affinis ZS10
    |    |--P. agavis M96
    |    |--P. anestios Williams 1985 GML13
    |    |--P. calceolariae CGW91 [=Trechocorys calceolariae K08]
    |    |--P. chenopodii Williams 1985 GML13
    |    |--P. citri ZS10
    |    |--P. comstocki M47
    |    |--P. crotonis G17
    |    |--P. elisae M96
    |    |--P. epidendrus Williams 1985 GML13
    |    |--P. eremosus Williams 1985 GML13
    |    |--P. gahani A71
    |    |--P. maritima A71
    |    |--P. markharveyi Gullan in Gullan, Moir & Leng 2013 GML13
    |    |--P. microcirculus M96
    |    |--P. nipae F37
    |    |--P. njalensis BW64
    |    `--P. viburni M96
    |--Rhizoecus M96
    |    |--R. americanus M96
    |    |--R. apizacos M96
    |    |--R. associatus M96
    |    |--R. cyperalis M96
    |    |--R. disjunctus M96
    |    |--R. falcifer M96
    |    |--R. gracilis M96
    |    |--R. leucosomus M96
    |    |--R. macgregori M96
    |    |--R. menkei M96
    |    |--R. mexicanus M96
    |    |--R. neomexicanus M96
    |    |--R. neostangei M96
    |    |--R. ovatus M96
    |    |--R. pauciporus M96
    |    |--R. polyporus M96
    |    |--R. solani M96
    |    |--R. spinipes M96
    |    |--R. stangei M96
    |    `--R. tropicalis M96
    `--Phenacoccus LS96 [Phenacoccinae KP01]
         |--P. bahiae C01
         |--P. calcitectus Cockerell 1901 C01
         |--P. capensis M96
         |--P. defectus M96
         |--P. eremicus M96
         |--P. eschscholtziae M96
         |--P. franseriae M96
         |--P. gossypii M96
         |--P. gregosus M96
         |--P. helianthi M96
         |--P. herreni M99
         |--P. hurdi M96
         |--P. incomptus M96
         |--P. madeirensis TKC07
         |--P. manihoti LS96
         |--P. mexicanus M96
         |--P. parvus M96
         |--P. psidiarum M96
         |--P. solani M96
         |--P. solenopsis M96
         |--P. spiniferus Hempel 1901 H01
         `--P. yuccae [incl. P. yuccae var. barberi] C01

*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.

[A71] Askew, R. R. 1971. Parasitic Insects. Heinemann Educational Books: London.

[BW64] Booth, C., & J. M. Waterston. 1964. Calonectria rigidiuscula. C.M.I. Descriptions of Pathogenic Fungi and Bacteria 21.

[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).

[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.

[F37] Ford, C. 1937. Minutes of the 481st regular meeting of the Entomological Society of Washington. Proceedings of the Entomological Society of Washington 39 (4): 88–90.

[G17] Girault, A. A. 1917. New Javanese Hymenoptera. 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: 59–71).

[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.

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

[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.

[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.

[M47] Mayr, E. 1947. Ecological factors in speciation. Evolution 1: 263–288.

[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.

[M99] Moraes, G. J. de. 1999. Pest status of the cassava green mite in Brazil and strategies for its control. In: Needham, G. R., R. Mitchell, D. J. Horn & W. C. Welbourn (eds) Acarology IX vol. 2. Symposia pp. 287–291. Ohio Biological Survey: Columbus (Ohio).

[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.

[TKC07] Tixier, M.-S., S. Kreiter & B. Cheval. 2007. Vineyard colonisation by predaceous mites (Acari: Phytoseiidae) living in surrounding vegetation. A three year study in the south of France. 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. 475–487. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[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.

[ZS10] Zborowski, P., & R. Storey. 2010. A Field Guide to Insects in Australia 3^rd ed. Reed New Holland: Sydney.

Last updated: 8 February 2020.

Pentatomoidea

Asopus reciprocus syntype, from the Natural History Museum.

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

The Pentatomoidea include the stink bugs and related taxa. Many pentatomoids have an enlarged scutellum, which in extreme cases may cover the entire abdomen. Such examples include the Plataspididae, in which the hemelytra are much longer than the body but folding to fit under the scutellum at rest. 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). Most pentatomoids are plant-feeders but members of the pentatomoid subfamily Asopinae are predators, and have a thick rostrum associated with bucculae that are shorted to allow greater flexibility of the labium for predation. Other distinctive members of the group 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).

Synapomorphies (from Shcherbakov & Popov 2002): Antennae 5-segmented; only two lateral trichobothria per abdominal segment; genital aperture directed caudally; egg barrel-shaped with circular eclosion rent.

<==Pentatomoidea GE05
    |--+--Lestonia China 1955 [Lestoniidae] G75
    |  |    |--*L. haustorifera China 1955 G75
    |  |    `--L. grossi CGW91
    |  `--Plataspididae [Brachyplatidae, Coptosomatidae, Plataspidinae] G75
    |       |--Brachyplatys D14
    |       |    |--B. pacificus Wall. 1851 D14
    |       |    `--B. vanikorensis Boisduval 1835 B35
    |       |--Coptosoma S07
    |       |    |--C. ostensum S07
    |       |    `--C. scutellatum H01
    |       |--Ceratocoris RD77
    |       `--Elapheozygum RD77
    `--+--Cydnidae G75
       `--+--Phloeidae G75
          `--Pentatomidae G75
               |  i. s.: Thyanta pallidovirens WS04
               |         Erthesina fullo YS01, H01
               |         Rhaphigaster nebulosa (Poda 1761) H01, L-SSY04 [incl. R. grisea H01]
               |         Mormidea integella [=Euschistus (Paraschistus) integellus] R74
               |         Irochrotus caspius H01
               |         Phimodera collina H01
               |         Masthletinus abbreviatus H01
               |         Neottiglossa pusilla [incl. N. inflexa] H01
               |         Eusarcoris aeneus H01
               |         Rubiconia intermedia H01
               |         Peribalus vernalis H01
               |         Palomena prasina H01
               |         Picromerus bidens H01
               |         Commius elegans C91
               |         Dalsira bohndorffi RD77
               |         Chlorochroa ligata RD77
               |--Trincavellius [Discocephalinae] B27
               |--Pentatominae SS02
               `--Asopinae [Asopidae] SS02
                    |--Ealda minax Walk. 1867 D14
                    |--Zicrona caerulea G76, RD77
                    |--Stiretrus G75
                    |--Eocanthocona MC13
                    `--Asopus D01
                         |--A. frontalis (Walker 1867) [=Strachia frontalis] D01
                         |--A. hamatus (Walker 1867) [=Strachia hamata; incl. S. saturata Walker 1867] D01
                         `--A. reciprocus (Walker 1867) [=Strachia reciproca; incl. S. megaspila Walker 1867] D01

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
  Dismegistus SDT88
  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).

[D01] Distant, W. L. 1901. Rhynchotal notes.—VIII. Heteroptera: fam. Coreidae. Annals and Magazine of Natural History, series 7, 7: 6–22.

[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.

[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.

[G76] Gross, G. F. 1976. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera—Part II. 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.

[L-SSY04] López-Sebastián, E., J. Selfa & J. Ylla. 2004. Primeros datos del parasitismo de Ooencyrtus pityocampae (Mercet, 1921) sobre Graellsia isabelae (Graells, 1849) en condiciones de laboratorio. Graellsia 60 (1): 121–123.

[MC13] Majer, J. D., S. K. Callan, K. Edwards, N. R. Gunawardene & C. K. Taylor. 2013. Baseline survey of the terrestrial invertebrate fauna of Barrow Island. Records of the Western Australian Museum Supplement 83: 13–112.

[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.

[R74] Rolston, L. H. 1974. Revision of the genus Euschistus in Middle America (Hemiptera, Pentatomidae, Pentatomini). Entomologica Americana 48 (1): 1–102.

[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.

[WS04] Wang, Q., & G. L. Shi. 2004. Mating frequency, duration, and circadian mating rhythm of New Zealand wheat bug Nysius huttoni White (Heteroptera: Lygaeidae). New Zealand Entomologist 27: 113–117.

[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: 8 February 2020.