Diagnostic features of Cymatopsocus opalinus, from New (1975).

Belongs within: Paraneoptera.
Contains: Psocathropetae, Trogioidea, Lepidopsocidae, Psocomorpha, Nanopsocetae, Amphientominae.

The Psocodea include the barklice, small insects that mostly feed on fungi and other micro-organisms, as well as the true parasitic lice. It is now generally accepted that the latter are a derived subgroup of the former. The most basal division in the Psocodea is between the clades Trogiomorpha and Psocina; Trogiomorpha have antennae with more than twenty segments and a strong posterior spine on the paraproct (the lateral sclerites alongside the anus) whereas Psocina have fewer antennal segments and lack the paraproct spine. Psocina also have the hypopharyngeal filaments fused for at least part of their length, while they remain separate in Trogiomorpha. Within the Psocina, members of the Troctomorpha lineage have antennae with at least some segments secondarily annulated, wings lacking a pterostigma and often a triangular or T-shaped sclerite on the large subgenital plate (Mockford 1993).

Characters (from Rasnitsyn 2002, for Psocidea): Size small, rarely medium. General appearance highly variable depending on free living or parasitic habit and mode of parasitism. Head with clypeus strongly convex to house cibarial muscles (secondarily less developed in some parasitic forms). Mandible chewing (reverted and thus tearing instead of biting in elephant louse, reduced in sucking lice). Lacinia free of remaining maxilla, rod-like. Hypopharynx with two ovoid sclerites. Pronotum small. Wings often lost, when present roof-like at rest, rather poor in venation, coupled in flight except in some Palaeozoic species with homonomous wings. Legs cursorial or, in parasites, clinging, with tarsus l-3-segmented (4-segmented in some extinct forms). 1st abdominal sternum lost. Ovipositor present or lost. Aedeagus lost in living species. Cercus lost. Extant forms with abdominal ganglia fused in single ganglionic mass and with 4 or less Malpighian tubes. Ovaries polytrophic. Chromosomes holokinetic.

Psocodea [Copeognatha, Corrodentia, Psocida, Psocidea, Psocoptera]
    |--Trogiomorpha [Trogiina] Y02
    |    |--Psocathropetae M93
    |    `--Atropetae [Trogiformia] L02
    |         |--Trogioidea R02
    |         `--Lepidopsocidae L02
    `--Psocina R02
         |--Psocomorpha Y02
         `--Troctomorpha [Amphientomiformia, Amphientomomorpha, Troctopsocoidea] Y02
              |--Nanopsocetae R02
              `--Amphientometae NL07
                   |  i. s.: Protroctopsocidae [Protroctopsocinae] NL07
                   |           |--Chelyopsocus Lienhard 1980 NL07, L98
                   |           |    `--*C. garganicus Lienhard 1980 L98
                   |           `--Philedaphia Lienhard 1995 NL07, L98
                   |                |--*P. aphrodite Lienhard 1995 [=Chelyopsocus aphrodite] L98
                   |                `--P. hauseri (Lienhard 1988) [=Chelyopsocus hauseri] L98
                   |         Troctopsocidae [Plaumanniidae, Troctopsocinae] NL07
                   |           |--*Selenopsocus schwendingeri Lienhard & Mockford 1997 NL07
                   |           |--Troctopsoculus NL07
                   |           |--Coleotroctellus Lienhard 1988 L02 [incl. Sinitroctopsocus Li 1993 NL07, L02]
                   |           |    |--*C. burckhardti Lienhard 1988 NL07, L02
                   |           |    |--C. loebli Lienhard 1988 NL07
                   |           |    |--C. sui (Li 1993) L02 [=*Sinitroctopsocus sui NL07]
                   |           |    `--C. venosus NL07
                   |           `--Thaipsocus NL07
                   |                |--*T. siamensis Lienhard & Mockford 1997 NL07
                   |                |--T. bau Lienhard & Mockford 1997 NL07
                   |                |--T. borneensis Lienhard & Mockford 1997 NL07
                   |                |--T. doisuthep Lienhard & Mockford 1997 NL07
                   |                |--T. orientalis (Lienahard 1990) NL07
                   |                |--T. sarawakensis Lienhard & Mockford 1997 NL07
                   |                `--T. thamluang Lienhard & Mockford 1997 NL07
                   |--Electrentomoidea R02
                   |    |  i. s.: Parelectrentomum S65
                   |    |--Musapsocidae [Musapsocoidea] NL07
                   |    |    |--*Musapsocus huastecanus NL07
                   |    |    `--*Musapsocoides nadleri NL07
                   |    `--Electrentomidae [Electrentominae, Manicapsocidae] NL07
                   |         |--Electrentomum S65
                   |         |--*Epitroctes tuxtlarum NL07
                   |         |--*Nothoentomum palpale NL07
                   |         |--*Phallopsocus carminatus NL07
                   |         `--Manicapsocus Smithers 1965 NL07, S65
                   |              `--*M. alettae Smithers 1965 S65
                   `--Amphientomoidea L02
                        |--Compsocidae L02
                        |    |--*Compsocus elegans NL07
                        |    `--*Electrentomopsis variegata NL07
                        `--Amphientomidae N75
                             |  i. s.: Neoseopsis Badonnel 1986 L02
                             |           `--*N. nuda Badonnel 1986 B86
                             |         Seopsocus Roesler 1940 T13
                             |--Amphientominae L02
                             |--Antivulgaris Li 2002 [Antivulgarinae] L02
                             |    `--*A. jianfenglingensis Li 2002 L02
                             |--Compressionis Li 1993 [Compressioninae] L02
                             |    `--*C. introvenis Li 1993 L02
                             `--Tineomorphinae N75
                                  |--Tineomorpha Enderlein 1906 N75
                                  |    |--*T. greeniana Enderlein 1906 NL07, N75
                                  |    |--T. angolana Badonnel 1955 N75
                                  |    |--T. jacobsoniana Enderlein 1926 N75
                                  |    |--T. murphyi New 1975 N75
                                  |    `--T. timahensis New 1975 NL07
                                  `--Cymatopsocus Enderlein 1903 T13
                                       |--*C. opalinus Enderlein 1903 N75
                                       |--C. emarginatus New 1975 N75
                                       |--C. enderleini New 1975 N75
                                       |--C. enigmaticus New 1975 N75
                                       `--C. maculatus Lee 1974 N75

Psocodea incertae sedis:
  Tshekarcephalus [Tshekarcephalidae] R02
    `--T. bigladipotens R02
  Surijokopsocus R02
  Sinopsocus oligovenus Lin 1976 L02
  Liaoxipsocus beipiaoensis (Hong 1983) L02
  Beipiaopsocus divaricatus (Hong 1983) L02
  Huaxiapsocus parvus (Hong 1983) L02
  Psocites pectinatus (Hong 1992) L02
  Diploperipsocus phagococcus Li & Mockford 1993 L02
  Mascaropsocus spinosus Badonnel & Pearman 1971 SN08
  Miotroctes Pierce 1960 YL10
    `--M. rousei Pierce 1960 YL10
  Westphalopsocus Azar, Nel et al. in Nel, Roques et al. 2013 [Westphalopsocidae] NR13
    `--*W. pumilio Azar, Nel et al. in Nel, Roques et al. 2013 NR13
  Zygopsocus [Zygopsocidae] NR13

*Type species of generic name indicated


[B86] Badonnel, A. 1986. Psocoptères de Colombie (Insecta, Psocoptera). Spixiana 9 (2): 179–223.

[L02] Li F. 2002. Psocoptera of China vol. 1. Science Press: Beijing.

[L98] Lienhard, C. 1998. Faune de France. France et Régions Limitrophes. 83. Pscoptères Euro-Méditerranéens. Fédération Française des Sociétés de Sciences Naturelles: Paris.

[M93] Mockford, E. L. 1993. North American Psocoptera (Insecta). Sandhill Crane Press, Inc.

[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.

[N75] New, T. R. 1975. Lepidopsocidae and Amphientomidae (Psocoptera) from Malaysia and Singapore. Oriental Insects 9 (2): 177–194.

[NL07] New, T. R., & C. Lienhard. 2007. The Psocoptera of Tropical South-east Asia. Brill: Leiden.

[R02] Rasnitsyn, A. P. 2002. Superorder Psocidea Leach, 1815. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 125–133. Kluwer Academic Publishers: Dordrecht.

[SN08] Schmidt, E. R., & T. R. New. 2008. The Psocoptera (Insecta) of Tasmania, Australia. Memoirs of Museum Victoria 65: 71–152.

[S65] Smithers, C. N. 1965. Descriptions and new records of Atropetae, Psocatropetae and Amphientometae (Psocoptera) from Africa. Journal Ent. Soc. S. Africa 28 (1): 44–49.

[T13] Taylor, C. K. 2013. The genus Lithoseopsis (Psocodea: Amphientomidae) in the Western Australian fauna, with description of the male of Lithoseopsis humphreysi from Barrow Island. Records of the Western Australian Museum Supplement 83: 245–252.

[Y02] Yoshizawa, K. 2002. Phylogeny and higher classification of suborder Psocomorpha (Insecta: Psocodea: ‘Psocoptera’). Zoological Journal of the Linnean Society 136: 371–400.

[YL10] Yoshizawa, K., & C. Lienhard. 2010. In search of the sister group of the true lice: a systematic review of booklice and their relatives, with an updated checklist of Liposcelididae (Insecta: Psocodea). Arthropod Systematics and Phylogeny 68 (2): 181–195.

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