Phthiraptera

Laemobothrion tinnunculi, copyright Museum of New Zealand Te Papa Tongarewa.

Belongs within: Nanopsocetae.
Contains: Philopteridae, Trichodectidae, Anoplura, Menoponidae.

The Phthiraptera, true lice, are a group of small, wingless insects that live as external parasites on warm-blooded vertebrates.

Synapomorphies (from Grimaldi & Engel 2005): Apterous; head with very limited movement; eyes highly reduced (only several facets), usually completely lost; ocelli absent; dorsal tentorial arms lost; lacinial stylets reduced or lost; maxillae reduced or lost; antennae reduced to three flagellomeres. Metathoracic spiracle absent; third thoracic ganglion fused with abdominal ganglion. Egg with hydropile and operculum, cemented to host hair or feather. Three nymphal instars.

<==Phthiraptera [Mallophaga, Nyrmida, Parasita, Pediculida, Philopterina, Siphunculata]
    |--Euphthiraptera [Ischnocera, Philopteroidea] A99
    |    |  i. s.: Syrrhaptoecus Waterson 1928 T62
    |    |         Fulicoffula lurida A71
    |    |         Rallicola A71
    |    |         Sturnidoecus sturni A71
    |    |         Dasyonygidae RD77
    |    |--+--Trichophilopterus RD77 [Trichophilopteridae GE05]
    |    |  |--Philopteridae GE05
    |    |  `--+--Goniodes CM91 [Goniodidae GE05]
    |    |     |    `--G. dissimilis CM91
    |    |     `--Rhopaloceras [Heptapsogasteridae] GE05
    |    |          `--R. rudimentarius GE05
    |    `--+--Trichodectidae GE05
    |       |--Saurodectes [Saurodectidae] R02
    |       |    `--S. vrsanskyi GE05
    |       `--Lipognatha A99
    |            |--Anoplura R02
    |            `--Haematomyzus CM91 [Haematomyzidae, Haematomyzoidea, Rhynchophthiraptera, Rhynchophthirina R02]
    |                 |--H. elephantis GE05
    |                 |--H. hopkinsi GE05
    |                 `--H. porci GE05
    `--Amblycera [Ricinoidea] GE05
         |  i. s.: Neomenopon Bedford 1920 T62
         |         Bonomiella Conci 1937 T62
         |         Piagetiella australis CM91
         |--Abrocomophagidae GE05
         |--Menoponidae CM91
         |--Trimenoponidae CM91
         |    |--Trimenopon hispidum (Burmeister 1838) T62
         |    `--Harrisonia uncinata GE05
         |--Gyropidae CM91
         |    |--Gliricola porcelli CM91
         |    `--Gyropus CM91
         |         |--G. gracilis G20 [incl. Pediculus porcelli Linnaeus 1758 G20, L58]
         |         `--G. ovalis CM91
         |--Ricinidae CM91
         |    |--Trochilocoetes RD77
         |    `--Ricinus CM91
         |         |--R. corvi (Linnaeus 1758) L02b, L58 [=Pediculus corvi L02b]
         |         |--R. diffusus RD77
         |         |--R. ernstlangi CM91
         |         `--R. pavonis (Linnaeus 1758) L02a, L58 [=Pediculus pavonis L02a]
         |--Laemobothrion [Laemobothriidae] CM91
         |    |--L. africanum Kellogg 1908 K08
         |    |--L. atrum CM91
         |    |--L. circi A71
         |    |--L. gypsis K08
         |    |--L. setigerum K08
         |    |--L. tinnunculi CM91
         |    `--L. titan K08
         `--Boopiidae CM91
              |--Paraheterodoxus insignis CM91
              |--Therodoxus CM91
              |--Boopia [incl. Phacogalia] C74
              `--Heterodoxus CM91
                   |--H. ampullatus CM91
                   |--H. longitarsus C70
                   |--H. octoseriatus CM91
                   `--H. spiniger CM91

Phthiraptera incertae sedis:
  Esthiopterum Harrison 1916 T62
    |--*E. gruis T62
    `--E. raphidium A71
  Pseudolipeurus plumbeus (Pessoa & Guimarães 1935) [incl. P. uoe Eichler 1950] T62
  Trogoninirmus Eichler 1944 T62
  Trogoniella Tendeiro 1960 T62
  Splendoroffula Clay & Meinertzhagen 1941 T62
  Falcolipeurus A71
  Trochiloecetes A71
  Aquanirmus A71
    |--A. americanus A71
    |--A. bucomfistii A71
    `--A. colymbinus A71
  Ibidoecus bisignatus A71
  Ferribia A71
  Campanulotes defectus GE05
  Neotrichodectes mephitidis HN90
  Bruelia S69
    |--B. saltatora Carriker 1956 S69
    `--B. violacea Carriker 1963 S69
  Heptapsus nothocercae Carriker 1936 S69
  Oxylipeurus S69
    |--O. craxae Carriker 1944 S69
    `--O. pauxus Carriker 1944 S69
  Amyrisidea simplex S69
    |--A. s. simplex S69
    `--A. s. pauxis Carriker 1950 S69
  Conciella clamatori Carriker 1966 S69
  Trochiliphagus mellivorus Carriker 1960 S69
  Strigiphilus E61
    |--S. acutifrons Emerson 1961 E61
    |--S. cursor (Burmeister 1838) E61
    `--S. oculatus (Rudow 1870) [incl. S. bubonis (Osborn 1896)] E61
  Kurodaia E61
    |--K. edwardsi Emerson 1961 E61
    |--K. keleri Emerson 1961 E61
    |--K. magna Emerson 1960 E61
    `--K. pectinata (Osborn 1902) E61
  Liotheidae K08
    |--Liotheum cornicis [=Ricinus cornicis] G20
    `--Trinoton K08
         |--T. femoratum Piaget 1880 PJ02
         `--T. luridum K08
  ‘Neumannia’ Uchida 1926 nec Trouessart 1888 nec Weise 1907 EH19

*Type species of generic name indicated

REFERENCES

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

[A99] Ax, P. 1999. Das System der Metazoa II. Ein Lehrbuch der phylogenetischen Systematik. Gustav Fisher Verlag: Stuttgart (translated: 2000. Multicellular Animals: The phylogenetic system of the Metazoa vol. 2. Springer).

[C70] Calaby, J. H. 1970. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 376–386. Melbourne University Press.

[C74] Calaby, J. H. 1974. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 51–52. Melbourne University Press.

[CM91] Calaby, J. H., & M. D. Murray. 1991. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 421–428. Melbourne University Press: Carlton (Victoria).

[E61] Emerson, K. C. 1961. Three new species of Mallophaga from the great horned owl. Proceedings of the Biological Society of Washington 74: 187–192.

[EH19] Énay, R., & M. K. Howarth. 2019. Part L, revised, volume 3B, chapter 7: Systematic descriptions of the Perisphinctoidea. Treatise Online 120: 1–184.

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[HN90] Hafner, M. S., & S. A. Nadler. 1990. Cospeciation in host-parasite assemblages: comparative analysis of rates of evolution and timing of cospeciation events. Systematic Zoology 39 (3): 192–204.

[K08] Kellogg, V. L. 1908. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 vol. 3 section 15. Corrodentia pt 4. Mallophaga. Almqvist & Wiksells Boktryckeri-A. B.: Uppsala.

[L02a] Latreille, P. A. 1802a. Histoire Naturelle des Fourmis, et recueil de mémoires et d'observations sur les abeilles, les araignées, les faucheurs, et autres insectes. Théophile Barrois père: Paris.

[L02b] Latreille, P. A. 1802b. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[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.

[PJ02] Palma, R. L., A. R. Johnson, F. Cezilly, F. Thomas & F. Renard. 2002. Diversity and distribution of feather lice on greater flamingoes (Phoenicopterus ruber roseus) in the Camargue, sothern France. New Zealand Entomologist 25: 87–89.

[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.

[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.

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

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

Trichodectidae

Geomydoecus costaricensis, from the Field Museum.

Belongs within: Phthiraptera.

The Trichodectidae are a group of lice found on mammals, characterised by a broadly oval body in both sexes, and short, thick legs with distally broadened tibiae, a terminal process and single claw.

<==Trichodectidae [Bovicolidae]
    |--Lorisicola GE05
    |--Werneckiella equi CM91
    |--Felicola subrostratus CM91
    |--Stachiella RD77
    |--Thomomydoecus Price & Emerson 1972 GE05, BP04
    |    |--T. barbarae GE05
    |    |--T. minor GE05
    |    `--T. wardii HN90
    |--Bovicola R02
    |    |--B. bovis CM91
    |    |--B. caprae Gurlt 1843 CM91, T62 [=Damalinia caprae T62]
    |    |--B. crassipes R02
    |    `--B. ovis CM91
    |--Damalinia C70
    |    |--D. americana A71
    |    |--D. bovis H74
    |    |--D. equi C70
    |    |--D. limbata (Gervais 1844) [=Bovicola limbatus] T62
    |    `--D. ovis H74
    |--Trichodectes GE05
    |    |--T. canis CM91 [=Ricinus canis G20]
    |    |--T. colobi Kellogg 1908 K08
    |    |--T. inaequalis K08
    |    |--T. latus G20
    |    |--T. melis A71
    |    `--T. octomaculatus GE05
    `--Geomydoecus Ewing 1929 GE05, BP04
         |--+--G. perotensis GE05
         |  `--G. thomomyus GE05
         `--+--+--G. chapini GE05
            |  `--+--+--G. cherriei GE05
            |     |  `--G. costaricensis GE05
            |     `--+--G. panamensis GE05
            |        `--G. setzeri GE05
            `--+--+--G. nadleri GE05
               |  `--G. trichopi GE05
               `--+--G. expansus GE05
                  `--+--+--G. geomydis GE05
                     |  `--G. oklahomensis GE05
                     `--+--G. actuosi GE05
                        `--+--G. ewingi GE05
                           `--G. texanus GE05

*Type species of generic name indicated

REFERENCES

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

[BP04] Banks, J. C., & A. M. Paterson. 2004. A penguin-chewing louse (Insecta: Phthiraptera) phylogeny derived from morphology. Invertebrate Systematics 18: 89–100.

[C70] Calaby, J. H. 1970. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 376–386. Melbourne University Press.

[CM91] Calaby, J. H., & M. D. Murray. 1991. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 421–428. Melbourne University Press: Carlton (Victoria).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[HN90] Hafner, M. S. & S. A. Nadler. 1990. Cospeciation in host-parasite assemblages: comparative analysis of rates of evolution and timing of cospeciation events. Systematic Zoology 39 (3): 192–204.

[H74] Helson, G. A. H. 1974. Insect Pests: Identification, life history, and control of pests of farms, horticulture, gardens, and public health. A. R. Shearer, Government Printer: Wellington (New Zealand).

[K08] Kellogg, V. L. 1908. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 vol. 3 section 15. Corrodentia pt 4. Mallophaga. Almqvist & Wiksells Boktryckeri-A. B.: Uppsala.

[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.

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

Philopteridae

Saemundssonia lari, from the Spencer Entomological Collection.

Belongs within: Phthiraptera.
Contains: Columbicola, Austrogoniodes.

The Philopteridae are a group of lice parasitic on birds, characterised by paired claws on the legs, a smoothly curved front margin on the clypeus, and a cylindrical last antennal segment (Richards & Davies 1977).

<==Philopteridae
    |--Saemundssonia Timmermann 1936 GE05, HPP80
    |    |--S. africana CM91
    |    |--S. calva A71
    |    |--S. celidoxa A71
    |    |--S. cephalus (Denny 1842) (see below for synonymy) P04
    |    |--S. desolata Timmermann 1959 HPP80
    |    |--S. fraterculae A71
    |    |--S. gaini (Neumann 1913) HPP80
    |    |--S. grylle A71
    |    |--S. lari (Fabricius 1780) HPP80
    |    |--S. lockleyi Clay 1949 HPP80
    |    |--S. merguli A71
    |    |--S. platygaster CM91
    |    |--S. stammeri Timmermann 1959 HPP80
    |    `--S. stresemanni Timmermann 1949 HPP80
    `--+--Austrogoniodes GE05
       `--+--Trabeculus Rudow 1866 GE05, HPP80
          |    |--T. flemingi GE05
          |    `--+--T. hexakon (Waterston 1914) GE05, HPP80
          |       `--T. schillingi Rudow 1866 GE05, HPP80
          `--+--Harrisoniella GE05
             |    |--H. densa RD77
             |    `--H. hopkinsi GE05
             `--Halipeurus Thompson 1936 GE05, HPP80
                  |--H. consimilis Timmermann 1960 GE05, HPP80
                  `--+--H. pelagicus GE05
                     `--+--H. falsus GE05
                        |    |--H. f. falsus HPP80
                        |    `--H. f. pacificus Edwards 1961 HPP80
                        `--+--H. diversus (Kellogg 1896) GE05, HPP80
                           `--H. spadix GE05

Philopteridae incertae sedis:
  Anatoecus CM91
    |--A. dentatus CM91
    `--A. pygaspis (Nitzsch in Giebel 1866) PJ02
  Megalipeurus von Kéler 1958 T62
    `--M. unicolor (Piaget 1880) (see below for synonymy) T62
  Anaticola Clay 1935 HPP80
    |--A. crassicornis (Scopoli 1763) HPP80
    |--A. ewingi (Qadri 1935) [=Columbicola ewingi] T62
    `--A. phoenicopteri (Coinde 1859) PJ02
  Turturicola Clay & Meinertzhagen 1937 T62
    |--*T. salimalii Clay & Meinertzhagen 1937 T62
    `--T. sudanicus (Mjöberg 1910) [=Lipeurus sudanicus, Columbicola sudanicus] T62
  Philopterus Nitzsch 1818 HPP80
    |--P. atratus A71
    |--P. communis [incl. Ricinus emberizae] G20
    |--P. corvi A71
    |--P. guttatus A71
    |--P. ocellatus A71
    `--P. turdi (Denny 1842) HPP80
  Nesiotinus Kellogg 1903 BP04
    `--N. demersus Kellogg 1903 BP04
  Columbicola T62
  Docophorus K08
    |--D. episcopi Kellogg 1908 K08
    |--D. icterodes K08
    |--D. indicus K08
    |--D. platystomus K08
    |--D. sphenophorus K08
    `--D. unifasciatus K08
  Nirmus K08
    |--N. furvus K08
    |--N. kilimandjarensis Kellogg 1908 K08
    |--N. opisthotomus Kellogg 1908 K08
    |--N. pileus K08
    `--N. signatus K08
  Lipeurus CM91
    |--L. afer Kellogg 1908 K08
    |--L. bifasciatus K08
    |--L. caponis CM91
    |--L. hebraeus K08
    |--L. heterographus RD77
    |--L. platalearum K08
    |--L. secretarius K08
    `--L. versicolor K08
  Brueelia Kéler 1936 HPP80
    |--B. cyclothorax (Burmeister 1838) HPP80
    `--B. merulensis (Denny 1842) HPP80
  Dahlemhornia CM91
  Cuclotogaster heterographus CM91
  Goniocotes Burmeister 1838 CM91, BP04
    |--G. bidentatus RD77
    `--G. gallinae CM91
  Degeeriella rufa RD77, T62
  Ardeicola Clay 1935 HPP80
    `--A. pilgrimi Tandan 1972 HPP80
  Docophoroides Giglioli 1864 HPP80
    |--D. brevis RD77
    |--D. harrisoni Waterston 1917 HPP80
    |--D. murphyi (Kellogg 1914) HPP80
    `--D. simplex (Waterston 1914) HPP80
  Haffneria Timmermann 1966 HPP80
    `--H. grandis (Piaget 1880) HPP80
  Naubates Bedford 1930 HPP80
    |--N. prioni (Enderlein 1908) HPP80
    `--N. pterodromi Bedford 1930 HPP80
  Paraclisis Timmermann 1965 HPP80
    |--P. diomedeae (Fabricius 1775) HPP80
    `--P. obscura (Rudow 1869) HPP80
  Pectinopygus Mjöberg 1910 HPP80
    |--P. dispar (Piaget 1880) HPP80
    `--P. gyricornis (Denny 1842) HPP80
  Pelmatocerandra Enderlein 1908 HPP80
    `--P. setosa (Giebel 1876) HPP80
  Perineus Harrison 1936 HPP80
    `--P. circumfasciatus Kéler 1957 HPP80
  Pseudonirmus Mjöberg 1910 HPP80
    `--P. gurli (Taschenberg 1882) HPP80
  Quadraceps Clay & Meinertzhagen 1939 HPP80
    |--Q. coenocoryphae Timmermann 1955 HPP80
    |--Q. punctatus (Burmeister 1838) HPP80
    `--Q. sellatus (Burmeister 1838) HPP80

Megalipeurus unicolor (Piaget 1880) [=Lipeurus unicolor, Oxylipeurus unicolor; incl. Lipeurus piageti Taschenberg 1882, Columbicola longiceps piageti] T62

Saemundssonia cephalus (Denny 1842) [incl. Docophorus atlanticus Kellogg 1914, Philopterus atlanticus, Saemundssonia atlantica] P04

*Type species of generic name indicated

REFERENCES

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

[BP04] Banks, J. C., & A. M. Paterson. 2004. A penguin-chewing louse (Insecta: Phthiraptera) phylogeny derived from morphology. Invertebrate Systematics 18: 89–100.

[CM91] Calaby, J. H., & M. D. Murray. 1991. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 421–428. Melbourne University Press: Carlton (Victoria).

[G20] Goldfuss, G. A. 1820. Handbuch der Naturgeschichte vol. 3. Handbuch der Zoologie pt 1. Johann Leonhard Schrag: Nürnberg.

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

[HPP80] Horning, D. S., R. L. Palma & R. L. C. Pilgrim. 1980. The lice (Insecta: Phthiraptera) from the Snares Islands, New Zealand. National Museum of New Zealand Miscellaneous Series 3: 1–17.

[K08] Kellogg, V. L. 1908. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 vol. 3 section 15. Corrodentia pt 4. Mallophaga. Almqvist & Wiksells Boktryckeri-A. B.: Uppsala.

[P04] Palma, R. L. 2004. Designation of a lectotype for Docophorus atlanticus Kellogg, 1914 (Insecta: Phthiraptera: Philopteridae). Tuhinga 15: 13–16.

[PJ02] Palma, R. L., A. R. Johnson, F. Cezilly, F. Thomas & F. Renard. 2002. Diversity and distribution of feather lice on greater flamingoes (Phoenicopterus ruber roseus) in the Camargue, sothern France. New Zealand Entomologist 25: 87–89.

[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.

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

Menoponidae

Colpocephalum fregili, copyright Natural History Museum, London.

Belongs within: Phthiraptera.

The Menoponidae are a diverse group of lice parasitic on birds.

Characters (from Richards & Davies 1977): Antennae four- or five-segmented, with 2 coeloconic sensilla on fourth or fourth and fifth segments; maxillary palps four-segmented; labial palps present, with five distal setae; pro-, meso- and metanota separate; mid and hind legs each with two tarsal claws; six pairs of spiracles on third to eighth abdominal segments; first abdominal tergum not fused with metanotum.

<==Menoponidae GE05
    |--Megamenopon rasnitsyni GE05
    |--Dennyus Neumann 1906 GE05, BP04
    |--Menacanthus Neumann 1912 HPP80
    |    |--M. eurysternus (Burmeister 1838) HPP80
    |    `--M. stramineus (Nitzsch 1818) [=Eomenacanthus stramineus] T62
    |--Kelerimenopon CM91
    |--Actornithophilus patellatus RD77, A71
    |--Austromenopon Bedford 1939 HPP80
    |    |--A. atrofulvum (Piaget 1880) HPP80
    |    |--A. bulleri Price & Clay 1972 HPP80
    |    |--A. elliotti Timmermann 1954 HPP80
    |    |--A. lutescens A71
    |    |--A. navigans (Kellogg 1896) HPP80
    |    |--A. paululum (Kellogg & Chapman 1899) HPP80
    |    |--A. pinguis (Kellogg 1896) HPP80
    |    `--A. stammeri Timmermann 1963 HPP80
    |--Myrsidea cucularis RD77
    |--Hohorstiella Eichler 1940 N91, T62
    |    `--H. lata N91
    |--Menopon CM91
    |    |--M. bucerotis Kellogg 1908 K08
    |    |--M. crocatum K08
    |    |--M. eulasius Kellogg 1908 K08
    |    |--M. forcipatum K08
    |    |--M. gallinae CM91
    |    |--M. pallidum RD77
    |    |--M. plegadis A71
    |    `--M. titan K08
    |--Ancistrona Westwood 1874 HPP80
    |--Longimenopon Thompson 1948 HPP80
    `--Colpocephalum CM91
         |--C. eucarenum K08
         |--C. extraneum K08
         |--C. falconii Carriker 1963 S69
         |--C. fregili CM91
         |--C. gracile K08
         |--C. grandiceps K08
         |--C. heterosoma Piaget 1880 PJ02
         |--C. miandrium Kellogg 1908 K08
         |--C. oreas Kellogg 1908 K08
         |--C. salimalii Clay 1951 PJ02
         |--C. semicinctum K08
         |--C. sjoestedti Kellogg 1908 K08
         |--C. subaequale K08
         |--C. trochioxum K08
         |--C. turbinatum CM91
         |--C. veratrum Kellogg 1908 K08
         `--C. xenicum Kellogg 1908 K08

*Type species of generic name indicated

REFERENCES

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

[BP04] Banks, J. C., & A. M. Paterson. 2004. A penguin-chewing louse (Insecta: Phthiraptera) phylogeny derived from morphology. Invertebrate Systematics 18: 89–100.

[CM91] Calaby, J. H., & M. D. Murray. 1991. Phthiraptera (lice). In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 421–428. Melbourne University Press: Carlton (Victoria).

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

[HPP80] Horning, D. S., R. L. Palma & R. L. C. Pilgrim. 1980. The lice (Insecta: Phthiraptera) from the Snares Islands, New Zealand. National Museum of New Zealand Miscellaneous Series 3: 1–17.

[K08] Kellogg, V. L. 1908. Wissenschaftliche Ergebnisse der Schwedischen Zoologischen Expedition nach dem Kilimandjaro, dem Meru und den Umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 vol. 3 section 15. Corrodentia pt 4. Mallophaga. Almqvist & Wiksells Boktryckeri-A. B.: Uppsala.

[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).

[PJ02] Palma, R. L., A. R. Johnson, F. Cezilly, F. Thomas & F. Renard. 2002. Diversity and distribution of feather lice on greater flamingoes (Phoenicopterus ruber roseus) in the Camargue, sothern France. New Zealand Entomologist 25: 87–89.

[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.

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

[T62] Tendeiro, J. 1962. Estudos sobre malófagos: Revisão monográfica do género Columbicola Ewing (Ischnocera, Philopteridae). Memórias da Junta de Investigações do Ultramar, ser. 2, 32: 7–460.

Psocathropetae

Psyllipsocus ramburii, copyright Chris Lann.

Belongs within: Psocodea.

The Psocathropetae are a group of barklice with a long head and long hind legs.

Characters (from New & Lienhard 2007): Head long, vertical. Antennae sometimes with secondary annulations on flagellar segments. Second segment of maxillary palp without conical sensillum on inner side. Fore wing with nodulus; brachypterous individuals with some venation discernible. Scales not present on body or wings. Combined length of hind tibia and tarsus greater than abdomen. Female with gonapophyses reduced but external valve present as broad setose lobe.

<==Psocathropetae [Psocatropetae, Psyllipsocetae, Psylliposciformia] NL07
    |--Prionoglarididae [Prionoglaretae, Prionoglaridae, Prionoglaroidea] NL07
    |    |  i. s.: *Sensitibilla strinatii NL07
    |    |--Speleketor Gurney 1943 M93 [Speleketoridae, Speleketorinae NL07, Speleketoroidea]
    |    |    |--*S. flocki Gurney 1943 M93
    |    |    |--S. irwini Mockford 1984 M93
    |    |    `--S. pictus Mockford 1984 M93
    |    `--Prionoglaridinae [Prionoglarinae] NL07
    |         |--*Siamoglaris zebrina Lienhard 2004 NL07
    |         `--Prionoglaris Enderlein 1909 [incl. Scoliopsyllopsis Enderlein 1912] G74
    |              |--*P. stygia Enderlein 1909 [incl. Scoliopsyllopsis latreillei Enderlein 1912] G74
    |              |--P. dactyloides Lienhard 1988 L98
    |              `--P. lindbergi Badonnel 1962 L98
    `--Psyllipsocidae [Psyllipsocinae, Psyllipsocoidea] NL07
         |--Pseudorypteryx García Aldrete 1984 M93
         |    `--*P. mexicana García Aldrete 1984 M93
         |--Pseudopsyllipsocus Li 2002 [=Pseudopsyllipsous] L03
         |    `--*P. gangliigerus Li 2002 [=P. gangliigera] L03
         |--Dorypteryx Aaron 1883 [incl. Dolopteryx Smithers 1958] M93
         |    |--*D. pallida Aaron 1883 [incl. D. albicans Ribaga 1907] M93
         |    |--D. domestica (Smithers 1958) [=*Dolopteryx domestica] LL09
         |    |--D. longipennis Smithers 1991 LL09
         |    `--D. yunnanica Li & Liu 2009 LL09
         |--Psocathropos Ribaga 1899 L02 (see below for synonymy)
         |    |--*P. lachlani Ribaga 1899 NL07 (see below for synonymy)
         |    |--P. cameriferus Li 2002 L02
         |    |--P. cuttackae [=*Granthakita cuttackae] NL07
         |    |--P. domesticus Li 2002 L03
         |    |--P. dyadoclemus Li 2002 L03
         |    |--P. pilipennis M93 [=*Gambrella pilipennis NL07]
         |    |--P. pleurotaenus Li 2002 L02
         |    |--P. purpuripsarous Li 2002 L02
         |    |--P. sinensis Li 1994 L02
         |    |--P. termitorum (n. d.) M93 [=*Vulturops termitorum NL07]
         |    `--P. tridymus Li 2002 L02
         `--Psyllipsocus Selys-Longchamps 1872 L02 (see below for synonymy)
              |--*P. ramburii Selys-Longchamps 1872 M93 (see below for synonymy)
              |--P. batuensis Thornton 1962 NL07
              |--P. hirsutus Thornton 1962 NL07
              |--P. maculatus Li 2002 L02
              |--P. metamicropterus (Enderlein 1963) L02
              |--P. oculatus Gurney 1943 M93
              |--P. sanxiaensis Li 1997 L02
              |--P. sauteri (Enderlein 1908) L02 [=*Parempheria sauteri NL07]
              |--P. sinicus Li & Yang 1988 L02
              |--P. spinosus Badonnel 1955 B55
              |--P. variabilis Badonnel 1986 B86
              |--P. yongi New & Lee 1992 NL07
              `--P. yucatan B50

Psocathropos Ribaga 1899 L02 [=Psocatropos NL07, Psocothropos (l. c.) L03; incl. Axinopsocus Enderlein 1903 M93, Gambrella Enderlein 1931 M93, Granthakita Behura & Dash 1968 L98, Psocinella Banks 1900 M93, Vulturops Townsend 1912 M93; Psocatropidae]

*Psocathropos lachlani Ribaga 1899 NL07 [incl. Vulturops floridensis Corbett & Hargreaves 1915 L98, V. floridanus (l. c.) L98, Psocatropos lesnei Badonnel 1931 M93, *Axinopsocus microps Enderlein 1903 NL07, L02, Psocathropos microps NL07, Psoquilla microps M93, *Psocinella slossonae Banks 1900 NL07, M93]

Psyllipsocus Selys-Longchamps 1872 L02 [incl. Fabrella Lacroix 1915 M93, Fita Navás 1913 M93, Nymphopsocus Enderlein 1903 M93, Ocellataria Weber 1907 M93, Ocelloria Weber 1906 M93, Parempheria Enderlein 1906 L98]

*Psyllipsocus ramburii Selys-Longchamps 1872 M93 [incl. Nymphopsocus troglodyta var. algericus Enderlein 1909 M93, Psy. ramburii brachypterus Badonnel 1943 M93, *Fabrella convexa Lacroix 1915 NL07, M93, *Nymphopsocus destructor Enderlein 1903 NL07, G74, *Ocelloria gravonymphia Weber 1906 NL07, L98, *Ocellatoria gravinympha NL07, Dorypteryx hageni Banks 1897 M93, Psocus pedicularius Rambur 1942 nec Hemerobius pedicularius Linnaeus 1758 nec P. pedicularius Burmeister 1839 M93, Nymphopsocus troglodyta Enderlein 1909 M93, Psyllipsocus (Nymphopsocus) troglodytes M93, Psy. ramburii troglodytes M93, *Fita vestigator Navás 1913 NL07, M93]

*Type species of generic name indicated

REFERENCES

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

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

[B50] Broadhead, E. 1950. A revision of the genus Liposcelis Motschulsky with notes on the position of this genus in the order Corrodentia and on the variability of ten Liposcelis species. Transactions of the Royal Entomological Society of London 101 (10): 335–388, pls 1–3.

[G74] Günther, K. K. 1974. Staubläuse, Psocoptera. Die Tierwelt Deutschlands 61: 1–315.

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

[LL09] Li, F. ,& X. Liu. 2009. Discovery in China of Dorypteryx Aaron (Psocoptera: Trogiomorpha: Psyllipsocidae), with one new species. Zootaxa 1983: 63–65.

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

[L03] Lienhard, C. 2003. Nomenclatural amendments concerning Chinese Psocoptera (Insecta), with remarks on species richness. Revue Suisse de Zoologie 110 (4): 695–721.

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

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

Lepidopsocidae

Fore (above) and hind wing of Nepticulomima orientalis, from New (1975).

Belongs within: Psocodea.
Contains: Echmepteryx.

The Lepidopsocidae are a group of barklice associated primarily with leaf litter and dead foliage (New & Lienhard 2007).

Characters (from New & Lienhard 2007): Body length usually ca 2 mm, fore wing length 2–2.5 mm. Lacinial apex divided with few teeth. Maxillary palpi with conical sensillum on inner side of second segment; labial palpi two-segmented. Antennae with numerous segments (more than 20, occasionally up to 50), not annulated. Tarsi three-segmented, claws curved, usually with one subapical tooth but sometimes with more or none. Fore wing often acuminate, usually (together with body and legs) with covering of flattened scales; pterostigma not thickened; vein Sc often well developed; R–Rs crossvein usually present; Rs and M commonly fused for a length; areola postica long; veins Cu₂ and 1A ending separately on wing margin. Hind wing vein R₁ long; vein M with two branches, these arising separately; hind wing sometimes absent. Paraproct with strong spine, trichobothrial field not defined. Female with gonapophyses reduced, ventral valve absent; dorsal valve small and often weakly sclerotised, sometimes absent; external valve large, setose. Male with phallosome with parameres divergent anteriorly; aedeagus often complex.

<==Lepidopsocidae [Perientometae, Perientomoidea]
    |--Parasoa [Parasoinae] NL07
    |    `--*P. haploneura Thornton 1962 NL07
    |--Lepolepis Enderlein 1906 L98 [Lepolepidinae NL07]
    |    |--*L. ceylonica Enderlein 1906 L98
    |    |--L. bicolor Broadhead 1955 L98
    |    `--L. columbiensis Badonnel 1986 B86
    |--Thylacellinae [Thylacellidae, Thylacelloidea, Udamolepinae] NL07
    |    |--Illepidopsocus Li 1994 L02
    |    |    `--*I. yunnanicus Li 1994 L02
    |    |--Pseudothylacus Li 2002 L02
    |    |    `--*P. acisopterus Li 2002 L02
    |    `--Thylacella Enderlein 1911 [incl. Udamolepis Enderlein 1912] M93
    |         |--*T. eversiana M93
    |         |--T. congolensis [=Udamolepis congolensis] B55
    |         |--T. cubana (Banks 1941) [=Echmepteryx cubana] M93
    |         |--T. fasciata Badonnel 1955 M93
    |         `--T. immaculata Badonnel 1955 B55
    |--Echinopsocinae NL07
    |    |--Scolopama NL07
    |    |--Pteroxanium Enderlein 1922 [incl. Lepidilla Ribaga 1905 non Matthew 1886, Tasmanopsocus Hickman 1934] M93
    |    |    `--*P. kelloggi (Ribaga 1905) (see below for synonymy) M93
    |    |--Echinopsocus NL07
    |    |    |--*E. erinaceus Enderlein 1903 NL07
    |    |    `--E. grayi NL07
    |    `--Neolepolepis Mockford 1993 M93
    |         |--*N. occidentalis (Mockford 1955) [=Lepolepis occidentalis] M93
    |         |--N. caribensis (Turner 1875) [=Lepolepsis (l. c.) caribensis] M93
    |         `--N. xerica [=Echmepteryx xerica] M93
    |--Lepidopsocinae NL07
    |    |--Echmepteryx NL07
    |    |--Cyptophania Banks 1931 NL07, MW13 [incl. Ptenocorium Enderlein 1931 MW13, Pteroxaniella Karny 1932 MW13]
    |    |    |--*C. hirsuta Banks 1931 MW13
    |    |    |--C. alutacea (Enderlein 1931) MW13 [=*Ptenocorium alutaceum NL07]
    |    |    |--C. australica Mockford & Wynne 2013 MW13
    |    |    |--C. bifurcata MW13 [=*Pteroxaniella bifurcata NL07]
    |    |    |--C. costalis Mockford & Wynne 2013 MW13
    |    |    |--C. marginata Thornton, Lee & Chui 1972 NL07
    |    |    `--C. pakaratii Mockford & Wynne 2013 MW13
    |    `--Lepidopsocus MW13
    |         |--*L. nepticulides Enderlein 1903 NL07
    |         |--L. cinctus MW13
    |         |--L. maculatus MW13
    |         |--L. magnus MW13
    |         |--L. marmoratus (Banks 1931) [=Echmepteryx marmorata] NL07
    |         |--L. pretiosus (Banks 1942) [=Echmepteryx pretiosus] NL07
    |         |--L. pseudomaculatus Thornton 1981 NL07
    |         `--L. sudarmani Vaughan, Thornton & New 1989 NL07
    `--Perientominae [Perientomidae] NL07
         |--*Papuapsocus caudatus Mockford 2005 NL07
         |--Proentomum Badonnel 1949 M93
         |    `--*P. personatum Badonnel 1949 M93
         |--Lepium Enderlein 1906 L02
         |    `--L. enderleini Banks 1937 L02
         |--Neolepipsocus Li 1992 L02
         |    `--*N. chinensis Li 1992 L02
         |--Perientomum NL07
         |    |--*P. trichopteryx [=Amphientomum trichopteryx] NL07
         |    `--P. corticola Vaughan, Thornton & New 1989 NL07
         |--Soa Enderlein 1904 M93
         |    |--*S. dahliana Enderlein 1904 NL07
         |    |--S. angolana Badonnel 1955 B55
         |    |--S. flaviterminata Enderlein 1906 NL07
         |    `--S. violacea M93
         `--Nepticulomima Enderlein 1906 M93
              |--*N. sakuntala Enderlein 1906 NL07
              |--N. biroiana (Enderlein 1903) NL07
              |--N. bothriata Thornton, Lee & Chui 1972 N75
              |--N. essigkeana Enderlein 1906 N75
              |--N. hoesemanni (Enderlein 1903) NL07 [=Perientomum hoesemanni S95]
              |--N. jacobsoni Enderlein 1926 NL07
              |--N. latisquama Enderlein 1926 NL07
              |--N. lusiae Thornton 1981 NL07
              |--N. orientalis New 1975 N75
              |--N. penicillata Enderlein 1926 NL07
              |--N. sumatrensis Vaughan, Thornton & New 1991 NL07
              `--N. uniformis Vaughan, Thornton & New 1989 NL07

*Pteroxanium kelloggi (Ribaga 1905) [=Lepidilla kelloggi; incl. Hyperetes brittanicus Harrison 1916, Tasmanopsocus litoralis Hickman 1934, Pteroxanium squamosum Enderlein 1922] M93

*Type species of generic name indicated

REFERENCES

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

[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. Psocoptères Euro-Méditerranéens. Fédération Française des Sociétés de Sciences Naturelles: Paris.

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

[MW13] Mockford, E. L., & J. J. Wynne. 2013. Genus Cyptophania Banks (Psocodea: 'Psocoptera': Lepidopsocidae): unique features, augmented description of the generotype, and descriptions of three new species. Zootaxa 3702 (5): 437–449.

[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.

[S95] Smithers, C. N. 1995. Psocoptera (Insecta) of Christmas Island. Invertebrate Taxonomy 9: 529–561.

Echmepteryx

Echmepteryx hageni, copyright Katja Schulz.

Belongs within: Lepidopsocidae.

Echmepteryx is a cosmopolitan genus of scaly barklice.

Characters (from Mockford 1993): Antenna with fewer than forty segments. Fore wing with vein R1 generally joined to Rs stem by crossvein; anterior wing catch relatively short and thick.

<==Echmepteryx Aaron 1886 M93
    |  i. s.: E. anomala S95
    |         E. frontalis Vaughan, Thornton & New 1991 NL07
    |         E. lacinipennis Enderlein 1926 NL07
    |         E. lawrencei Smithers 1995 NL07
    |         E. lineata Vaughan, Thornton & New 1991 NL07
    |         E. malayensis New 1975 NL07
    |         E. pletschi S95
    |         E. similis Badonnel 1955 B55
    |--E. (Echmepteryx) NL07
    |    |--*E. (E.) hageni (Packard 1870) M93 [=Amphientomum hageni NL07; incl. E. agilis Aaron 1886 M93]
    |    |--E. (E.) barba M93
    |    |--E. (E.) intermedia Mockford 1974 M93
    |    |--E. (E.) lealae M93
    |    |--E. (E.) submontana M93
    |    |--E. (E.) vara M93
    |    `--E. (E.) youngi Mockford 1974 M93
    |--E. (*Loxopholia) pinnula NL07
    |--E. (*Oxypsocus) hamiltoni NL07
    |--E. (*Thylacomorpha) symmetrolepis NL07
    `--E. (Thylacopsis Enderlein 1911) M93
         |--E. (*T.) mihira [=Thylax mihira] NL07
         |--E. (T.) falco (Badonnel 1949) M93 [=Thylacopsis falco B49]
         |--E. (T.) lunulata Thornton, Lee & Chui 1972 NL07
         |--E. (T.) madagascariensis (Kolbe 1885) NL07 (see below for synonymy)
         `--E. (T.) pallida Smithers 1965 S95

Echmepteryx (Thylacopsis) madagascariensis (Kolbe 1885) NL07 [=Thylax madagascariensis M93; incl. Thylacopsis albidus Badonnel 1949 M93, Echmepteryx albidus M93, E. costalis Banks 1931 M93, Lepidopsocus costalis M93]

*Type species of generic name indicated

REFERENCES

[B49] Badonnel, A. 1949. Psocoptères de la Cote d'Ivoire. Mission Paulian-Delamare (1945). Revue Française d'Entomologie 16: 20–46.

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

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

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

[S95] Smithers, C. N. 1995. Psocoptera (Insecta) of Christmas Island. Invertebrate Taxonomy 9: 529–561.

Lichenomiminae

Lichenomima sp., copyright Katja Schulz.

Belongs within: Psocetae.

The Lichenomiminae are a group of barklice of the Myopsocidae in which the hind wing has veins Rs and M joined by a crossvein (Li 2002).

<==Lichenomiminae L02
    |--Lophomyini L02
    |    |--Lophomyus Li 2002 L02
    |    |    |--*L. bidigitatus Li 2002 L02
    |    |    `--L. petersi (Smithers 1971) [=Lophopterygella petersi] L02
    |    `--Lophopterygella Enderlein 1907 [incl. Festona Navas 1922] L02
    |         |--*L. camelina Enderlein 1907 NL07
    |         |--L. bellula Li 2002 L02
    |         |--‘*Festona’ lunata NL07
    |         |--L. ridderi Vaughan, Thornton & New 1991 NL07
    |         |--L. spilota Smithers & Thornton 1979 NL07
    |         `--L. triangularis Lienhard 2004 NL07
    `--Lichenomimini L02
         |--Gyromyus Li 2002 L02
         |    |--*G. gyrus Li 2002 L02
         |    `--G. sanguensis (New 1973) [=Myopsocus sanguensis, Lichenomima sanguensis] L02
         `--Lichenomima Enderlein 1910 L02 [=Lachenomima (l. c.) L03, Licnenomima (l. c.) L03]
              |--*L. conspersa NL07
              |--L. ampla (Smithers & Thornton 1974) NL07
              |--L. camelina Enderlein 1907 L02
              |--L. chelata (Thornton & Woo 1973) L02
              |--L. clypeofasciata (Mockford 1974) L02
              |--L. coloradensis (Banks 1907) [=Myopsocus coloradensis] M93
              |--L. corniculata Li 1995 L02
              |--L. cubitalis Lienhard 2004 NL07
              |--L. cylindra Li 2002 L03
              |--L. elongata (Thornton 1960) [=Myopsocus elongatus] L02
              |--L. excavata Li 2002 L03
              |--L. fasciata Badonnel 1946 L02
              |--L. fenestrata Enderlein 1926 NL07
              |--L. gibbulosa Li 2002 L02
              |--L. guineensis Enderlein 1914 B49
              |--L. hamata Li 1995 L02
              |--L. hangzhouensis Li 2002 L02
              |--L. harpeodes Li 2002 L02
              |--L. intermedia Pearman 1934 L02 [=L. cameruna var. intermedia B55]
              |--L. leucospila Li 2002 L02
              |--L. lugens (Hagen 1861) (see below for synonymy) M93
              |--L. machadoi (Badonnel 1977) L02
              |--L. maxima (Smithers 1957) L02
              |--L. merapi Thorton in Thornton & Browning 1992 NL07
              |--L. muscosa Y02
              |--L. orbiculata Li 2002 L03
              |--L. oxycera Li 2002 L02
              |--L. pauliani Badonnel 1949 B55
              |--L. sinuosa Lienhard 2004 NL07
              |--L. sparsa (Hagen 1861) M93 [=Psocus sparsus M93, Myopsocus sparsus M74]
              |--L. sumatrana (Enderlein 1906) NL07
              |--L. tridens Li 2002 L02
              `--L. unicornis Li 2002 L02

Lichenomima lugens (Hagen 1861) [=Psocus lugens, Myopsocus lugens; incl. P. virginianus Banks 1900, M. virginianus] M93

*Type species of generic name indicated

REFERENCES

[B49] Badonnel, A. 1949. Psocoptères de la Cote d'Ivoire. Mission Paulian-Delamare (1945). Revue Française d'Entomologie 16: 20–46.

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

[L02] Li F. 2002. Psocoptera of China vol. 2. Science Press: Beijing.

[L03] Lienhard, C. 2003. Nomenclatural amendments concerning Chinese Psocoptera (Insecta), with remarks on species richness. Revue Suisse de Zoologie 110 (4): 695–721.

[M74] Mockford, E. L. 1974. Records and descriptions of Cuban Psocoptera. Entomologica Americana 48 (2): 103–215.

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

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

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

Myopsocinae

Myopsocus eatoni, copyright Katja Schulz.

Belongs within: Psocetae.

The Myopsocinae are a group of barklice of the Myopsocidae in which the hind wing has veins Rs and M fused for a length, and the outer edge of the fore wing is not curved (Li 2002).

<==Myopsocinae L02
    |--Smithersia Thornton in Thornton & Browning 1992 L03 (see below for synonymy)
    |    |--*S. hermosa (Banks 1920) [=Psocus hermosus] NL07
    |    |--S. angusta (Vaughan, Thornton & New 1991) NL07
    |    |--S. hermosoides (Vaughan, Thornton & New 1991) NL07
    |    |--S. newi Thornton in Thornton & Browning 1992 NL07
    |    |--S. scapiformis (Li 2002) [=Polygonomyus scapiformis] L03
    |    |--S. sexangula (Li 2002) [=Polygonomyus sexangulus] L03
    |    `--S. sinica (Li 2002) [=*Polygonomyus sinicus] L03
    `--Myopsocus Hagen 1866 [incl. Phlotodes Enderlein 1910, Rhaptoneura Enderlein 1910] L02
         |--*M. unduosus (Hagen 1859) [=Psocus unduosus] NL07
         |--‘Rhaptoneura’ africana Badonnel 1955 B55
         |--‘Phlotodes’ angolensis Badonnel 1955 B55
         |--M. angulatus Vaughan, Thornton & New 1991 NL07
         |--M. angustus S95
         |--M. antillanus (Mockford 1974) [=Phlotodes antillanus] M93
         |--M. apicalis (Thornton 1984) NL07
         |--‘Phlotodes’ australis GE05
         |--M. bakeri Banks 1916 NL07
         |--M. bomasa (Smithers & Thornton 1974) NL07
         |--M. brunneigena (Smithers & Thornton 1979) NL07
         |--M. brunneipes Smithers 1995 NL07
         |--M. cameruna Enderlein 1903 M74, B55 [=Lichenomima cameruna B55]
         |--M. clypeofasciatus Mockford 1974 M74
         |--‘Phlotodes’ congolensis B55
         |--M. corticola Vaughan, Thornton & New 1991 NL07
         |--‘Phlotodes’ cubanus Mockford 1974 M74
         |--M. dentatus (Smithers & Thornton 1974) NL07
         |--‘*Rhaptoneura’ dispar NL07
         |--M. eatoni McLachlan 1880 L98 [=Rhaptoneura eatoni M93; incl. M. eatoni var. europaeus Ribaga 1907 L98]
         |--M. enderleini Banks 1913 NL07
         |--M. fasciatus M74
         |--M. fenestratus (Smithers & Thornton 1974) NL07 [=Phlotodes fenestratus S95]
         |--M. gressitti (Smithers & Thornton 1974) NL07
         |--M. griseipennis S70
         |--M. hermosoides S95
         |--M. horripilis Lienhard 2004 NL07
         |--M. intermedia M74
         |--M. javensis Vaughan, Thornton & New 1991 NL07
         |--M. kolbei Enderlein 1903 [=*Phlotodes kolbei] NL07
         |--M. lineatus (Smithers & Thornton 1979) NL07
         |--M. lombokensis (Thornton 1984) NL07
         |--M. longigena (Enderlein 1926) NL07
         |--M. loriai Ribaga 1908 NL07
         |--M. luteolus (Banks 1920) NL07
         |--M. maculatus (Smithers & Thornton 1974) NL07
         |--M. marginatus (Smithers & Thornton 1974) NL07
         |--M. minutus (Mockford 1974) [=Phlotodes minutus] M93
         |--M. mjoebergi (Karny 1925) NL07
         |--M. palauensis Thornton, Lee & Chui 1972 L02
         |--M. pauliani M74
         |--M. peltatus (Smithers & Thornton 1974) NL07
         |--M. pilipes (Smithers & Thornton 1974) NL07
         |--M. platyvalvula (Smithers & Thornton 1979) NL07
         |--M. preclarus (Smithers & Thornton 1974) NL07 [=Phlotodes preclarus S95]
         |--M. pulchellus (Banks 1920) NL07
         |--M. rimosus (Smithers & Thornton 1974) NL07
         |--M. scabiosus (Smithers & Thornton 1974) NL07
         |--M. schoutedeni M74
         |--M. spatulatus Thornton in Thornton & Browning 1992 NL07
         |--M. thecatus (New & Thornton 1976) NL07
         |--M. toxeres (Smithers & Thornton 1974) NL07
         |--M. venustus (Smithers & Thornton 1974) NL07
         `--M. yunnanicus Li 2002 L02

Smithersia Thornton in Thornton & Browning 1992 L03 [incl. Polygonomyus Li 2002 L02, Ploygonomyus L02, Polyonomyus L03]

*Type species of generic name indicated

REFERENCES

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

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

[L02] Li F. 2002. Psocoptera of China vol. 2. Science Press: Beijing.

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

[L03] Lienhard, C. 2003. Nomenclatural amendments concerning Chinese Psocoptera (Insecta), with remarks on species richness. Revue Suisse de Zoologie 110 (4): 695–721.

[M74] Mockford, E. L. 1974. Records and descriptions of Cuban Psocoptera. Entomologica Americana 48 (2): 103–215.

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

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

[S70] Smithers, C. N. 1970. Psocoptera (psocids, booklice). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 367–375. Melbourne University Press.

[S95] Smithers, C. N. 1995. Psocoptera (Insecta) of Christmas Island. Invertebrate Taxonomy 9: 529–561.

Psocidus

Psocidus bifurcatus, from Li (2002).

Belongs within: Psocinae.

Psocidus is a genus of barklice historically used as a 'holding genus' for species of Psocini that could not be attributed to the restricted genus Psocus, but since redefined to include a smaller assemblage of species found in Africa and southern Asia (New & Lienhard 2007).

Characters (from New & Lienhard 2007): Male with hypandrium bearing setose apical lobe; phallosome with long anterior lateral arms, lateral apophyses, and pair of long inner sclerites; paraprocts bearing two trichobothrial fields, corresponding to main field divided by sensilla-free zone.

<==Psocidus Pearman 1934 L02 [=Psocodus (l. c.) L03; incl. Clistopsocus Navas 1924 L02, Sigmatina Enderlein 1925 L02]
    |--*P. zanzibarensis Pearman 1934 L02
    |--P. albovarius (Banks 1920) NL07
    |--P. apertus Thornton 1984 NL07
    |--P. arenosus (Navás 1922) NL07
    |--P. bifurcatus Li 2002 [=P. bicruris Li 2002 non Li 1990] L03
    |--P. biguttatus B55
    |--P. borneensis (Banks 1920) NL07
    |--P. divisus (Navás 1922) NL07
    |--P. femoratus (MacLachlan 1866) L02
    |--P. filicornis Enderlein 1903 L02
    |--P. gloriosus (Banks 1920) NL07
    |--P. guilinensis Li 2002 L02
    |--P. guttulatus Pearman 1934 B55
    |--P. hainanensis Li 2002 L02
    |--P. jacobsoni (Enderlein 1907) NL07
    |--P. javanicus (Enderlein 1907) NL07
    |--P. lacroixi (Navás 1920) NL07
    |--P. lestagei (Navás 1922) NL07
    |--P. lobatus Endang & Thornton 1992 NL07
    |--P. longicaudis B55
    |--P. luotongshanicus Li 2002 L02
    |--P. masinus (Navás 1923) NL07
    |--P. murphyi New 1975 NL07
    |--P. oxyurus Badonnel 1955 B55
    |--P. peregrans Vaughan, Thornton & New 1989 NL07
    |--P. proctus Vaughan, Thornton & New 1989 NL07
    |--P. reidi Thornton 1984 NL07
    |--P. relativus (Banks 1920) NL07
    |--P. samchiensis New 1978 L02
    |--P. sauteri (Enderlein 1908) L02
    |--P. segmentatus Endang & Thornton 1992 NL07
    |--P. similaris (Banks 1920) NL07
    |--P. strictus Thornton 1960 L02
    |--P. tacaoensis (Enderlein 1908) L02
    |--P. tergatus Endang & Thornton 1992 NL07
    |--P. validus Thornton 1960 NL07
    |--P. viscayanus (Banks 1920) NL07
    `--P. vitalisi (Navás 1922) NL07

*Type species of generic name indicated

REFERENCES

[B55] Badonnel, A. 1955. Psocoptères de l'Angola. Diamang Publicações Culturais 26: 11–267.

[L02] Li F. 2002. Psocoptera of China vol. 2. Science Press: Beijing.

[L03] Lienhard, C. 2003. Nomenclatural amendments concerning Chinese Psocoptera (Insecta), with remarks on species richness. Revue Suisse de Zoologie 110 (4): 695–721.

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