Fossil of Pincombea sp., from Shcherbakov & Popov (2002).

Belongs within: Hemiptera.
Contains: Aleyrodidae, Psylloidea, Coccinea, Aphidoidea.

The Sternorrhyncha are a group of small plant-sucking bugs including the aphids, scales, whiteflies and plantlice. They are commonly sedentary except when dispersing, moving little if at all. They are usually thought to be divided between two main lineages, of which the Psyllomorpha (including the whiteflies and plantlice) have jumping hind legs and eggs with short peduncles that may absorb water from the host plant (Grimaldi & Engel 2005).

The aphids (Aphidomorpha) and scale insects (Coccidomorpha) form a clade with polymorphic individuals within a population, reduced or absent male genitalia, and winged morphs with a composite vein in the forewing comprised of R and the bases of M and Cu (Grimaldi & Engel 2005). The first stem representatives of this clade (Aphidiformes) are the Permian Boreoscytidae, relatively large species with clinging legs that may have fed on large ovules (Shcherbakov & Popov 2002).

The fossil record of the Aphidomorpha goes back to the Triassic; the Late Triassic Creaphis theodora is a stem-aphid differing from later forms in having forewings with a long stem to vein Cu, and long straight Rs and M veins (Grimaldi & Engel 2005). The Naibiidae are a group of superficially aphid-like bugs known from the mid-Triassic to the early Palaeocene. Though lacking evidence of sexual dimorphism, they share a number of features with scale insects such as a dorsoventrally flattened head and few, widely-spaced ommatidia, and are likely stem-group relatives of the Coccinea (Shcherbakov & Popov 2002).

Characters (from Grimaldi & Engel 2005): Head extremely opisthognathous, with labium originating from or close to prosternum; proboscis with fine, hairlike feeding stylets, coiled in head capsule when at rest. Antenna with eight or fewer flagellomeres. Hind wing lacking vannus and vannal fold. Tarsus one- or two-segmented.

<==Sternorrhyncha [Gallinsecta, Phytadelga, Phytophthires]
    |  i. s.: Schizaphis graminumH06
    |--Psyllomorpha [Psyllina]GE05
    |    |--AleyrodidaeGE05
    |    `--PsylloideaGE05
    `--Aphidiformes [Aphidinea, Canadaphidoidea, Pincombeoidea, Pincombeomorpha]SLN15
         |  i. s.: Atheroides brevicornisK09
         |    |--Megaleurodes megocellata Hamilton 1990RJ93
         |    |--DinoscytaSLN15
         |    |--BoreoscytaSLN15
         |    `--ArchescytinopsisNR13
         `--+--+--Simulaphis [Simulaphididae]SLN15
            |  |    `--S. shaposhnikoviSLN15
            |  `--Pincombeidae [Pincombaeidae]SLN15
            |       |--Pincombea mirabilis Tillyard 1922F71
            |       `--Madygenopsyllidium djailautshoense Bekker-Migdisova 1985SLN15, RJ93
               `--Aphidomorpha [Aphidina, Gradipedes]SLN15
                    |  i. s.: Essigella californicaSC04
                    |         Dubiaphis curvataSLN15
                    |         ObtusicaudaPS96
                    |           |--O. coweniPS96
                    |           `--O. jonesiPS96
                    |         Liosomaphis berberidisPS96
                    |         KakimiaPS96
                    |           |--K. aquilegiaePS96
                    |           `--K. cynosbatiPS96
                    |         Eoessigia longicaudaPS96
                    |         Utamphorophora humboldtiPS96
                    |         Fimbriaphis wakibaePS96
                    |         Zyxaphis oregonensisPS96
                    |         Brachycorynella asparagiPS96
                    |         Hayhurstia atriplicisPS96
                    |         Aphthargelia symphoricarpiPS96
                    |         PterocommaPS96
                    |           |--P. bicolorPS96
                    |           `--P. flocculosaRD77
                    |         Cedoaphis incognitaPS96
                    |         Braggia eriogoniPS96
                    |         Nearctaphis clydesmithiPS96
                    |         Ceruraphis viburnicolaPS96
                    |         Monelliopsis caryaePS96
                    |         Oestlundiella flavaPS96
                    |         Dilachnus piceaeH44
                    |         Rasnitsynaphis Homan & Wegierek 2011 [Rasnitsynaphididae]HW11
                    |           |--*R. ennearticulata Homan & Wegierek 2011HW11
                    |           |--R. coniuncta Homan & Wegierek 2011HW11
                    |           `--R. quadrata Homan & Wegierek 2011HW11
                    |    |--SinojuraphididaeSLN15
                    |    |--DracaphididaeSLN15
                    |    `--NaibiidaeSLN15
                    |         |--NaibiaSLN15
                    |         |--CoccavusSLN15
                    |         `--PanirenaSLN15
                       |--GenaphisGE05 [Genaphididae, GenaphidoideaSLN15]
                       |    `--G. valdensisRJ93
                       |--Lutevanaphis Szwedo, Lapeyrie & Nel 2015 [Lutevanaphididae, Lutevanaphidoidea]SLN15
                       |    `--*L. permiana Szwedo, Lapeyrie & Nel 2015SLN15
                       |    |--Triassoaphis [Triassoaphididae]SLN15
                       |    |    `--T. cubitus Evans 1956P92
                       |    |--Creaphis [Creaphididae]SLN15
                       |    |    `--C. theodora Shcherbakov & Vengerek 1991RJ93
                       |    `--Leaphis [Leaphididae, Leaphidinae]SLN15
                       |         `--L. primaSLN15
                            |--SzelegiewicziaSP02 [SzelegiewicziidaeHW11, Szelegiewicziinae]
                            |    `--S. maculataSP02
                            |    |--Palaeoaphis archimediaRJ93
                            |    `--AmbaraphisP92
                            `--Ellinaphis Shaposhnikov 1979H02 [EllinaphididaeHW11]
                                 |--*E. sensoriata Shaposhnikov 1979H02
                                 `--E. leptoneura (Zhang et al. 1989) (see below for synonymy)H02

Ellinaphis leptoneura (Zhang et al. 1989) [=Caudaphis leptoneura; incl. C. minulissima Zhang et al. 1989, C. spinalis Zhang et al. 1989]H02

*Type species of generic name indicated


[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.

[H44] Hammer, M. 1944. Studies on the oribatids and collemboles of Greenland. Meddelelser om Grønland 141 (3): 1–210.

[H06] Hassanin, A. 2006. Phylogeny of Arthropoda inferred from mitochondrial sequences: strategies for limiting the misleading effects of multiple changes in pattern and rates of substitution. Molecular Phylogenetics and Evolution 38: 100–116.

[HW11] Homan, A., & P. Wegierek. 2011. A new family of aphids (Hemiptera, Sternorrhyncha) from the Lower Cretaceous of Baissa, Transbaikalia. ZooKeys 130: 167–174.

[H02] Hong Y. 2002. Amber Insect of China. Beijing Scientific and Technological Publishing House.

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

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[PS96] Pike, K. S., P. Starý, R. Miller, D. Allison, L. Boydston, G. Graf & T. Miller. 1996. New species and host records of aphid parasitoids (Hymenoptera: Braconidae: Aphidiinae) from the Pacific northwest, U.S.A. Proceedings of the Entomological Society of Washington 98 (3): 570–591.

[P92] Poinar, G. O., Jr. 1992. Life in Amber. Stanford University Press: Stanford.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th 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.

[SP02] Shcherbakov, D. E., & Yu. A. Popov. 2002. Superorder Cimicidea Laicharting, 1781: Order Hemiptera Linné, 1758. The bugs, cicadas, plantlice, scale insects, etc. [=Cimicida Laicharting, 1781, =Homoptera Leach, 1815 + Heteroptera Latreille, 1810). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 143–157. Kluwer Academic Publishers: Dordrecht.

[SC04] Stone, C., & N. C. Coops. 2004. Assessment and monitoring of damage from insects in Australian eucalypt forests and commercial plantations. Australian Journal of Entomology 43 (3): 283–292.

[SLN15] Szwedo, J., J. Lapeyrie & A. Nel. 2015. Rooting down the aphid’s tree—the oldest record of the Aphidomorpha lineage from Palaeozoic (Insecta: Hemiptera). Systematic Entomology 40: 207–213.

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