Psychodinae

Psychoda alternata, copyright Ivan Stankovic.

Belongs within: Psychodoidea.

The Psychodinae, moth flies, are a group of densely hairy flies whose larvae are most commonly found in moist or semi-aquatic habitats (Wagner & Ibáñez-Bernal 2009).

Characters (from Wagner & Ibáñez-Bernal 2009): Eye reniform with extension towards midline forming eye bridge above antennal insertions; eyes continuous or separated, rarely with eye bridge short and acutely pointed; flagellomeres fusiform, pyriform or nodiform.

<==Psychodinae WI-B09
    |--Maruina [Maruinini] WI-B09
    |    `--M. hirta WM96
    |--Pericomini WI-B09
    |    |--Lepidiella WI-B09
    |    `--Pericoma (Clytocerus) americana WI-B09, QV81
    |--Paramormiini WI-B09
    |    |--Clogmia albipunctata WI-B09, WT11
    |    |--Telmatoscopus mergacolis WI-B09
    |    `--Paramormia [incl. Duckhousiella] WI-B09
    |         |--P. furcata WI-B09
    |         `--P. ustulata WI-B09
    |--Mormiini WI-B09
    |    |--Atrichobrunettia alternata WI-B09, CM91
    |    |--Bryopharsos WI-B09
    |    |--Eugenys clavellata WI-B09
    |    |--Mormia O98
    |    `--Brunettia [incl. Neobrunettia Bravo & Amorim 1995] WI-B09
    |         `--B. dominicana WI-B09
    |--Psychodini WI-B09
    |    |--Trichopsychoda insulicola WI-B09, QV81
    |    |--Threticus WI-B09
    |    |--Feuerborniella WI-B09
    |    |--Philosepedon WI-B09
    |    |    |--P. helicis WM96
    |    |    |--P. mauroae Wagner & Masteller 1996 WM96
    |    |    |--P. opposita WM96
    |    |    `--P. quatei WM96
    |    `--Psychoda WI-B09
    |         |--P. alternata CM91
    |         |--P. cinerea WI-B09
    |         |--P. harrisi M83
    |         |--P. juliae Wagner & Masteller 1996 WM96
    |         |--P. ocellaris G20
    |         |--P. phalaenoides WI-B09
    |         |--P. puertoricana Wagner & Masteller 1996 WM96
    |         |--P. satchelli WM96
    |         `--P. spatulata M83
    `--Setomimini WI-B09
         |--Nemoneura liparotes WI-B09
         |--Didicrum pyramidon WI-B09
         |--Desmioza WI-B09
         |--Didimioza symphilia WI-B09
         |--Thrysocanthus WI-B09
         |--Valerianna WI-B09
         |--Neoarisemus maesi WI-B09
         |--Neurosystasis WI-B09
         |--Armillipora selvica Quate 1996 WI-B09
         |--*Australopericoma caudata [incl. A. wirthi] WI-B09
         |--Balbagathis discuspis WI-B09
         |--Setomima WI-B09
         |--Platyplastinx WI-B09
         |--Micrommatos WI-B09
         |    |--M. simplex WI-B09
         |    `--M. sylvaticum WI-B09
         |--Caenobrunettia WI-B09
         |    |--C. sarculosa WI-B09
         |    `--C. subditicia WI-B09
         |--Tonnoira WI-B09
         |    |--T. cavernicola WI-B09
         |    `--T. rapiformis WI-B09
         |--Alepia WI-B09
         |    |--A. diocula WI-B09
         |    |--A. hirtiventris WI-B09
         |    `--A. longinoi WI-B09
         `--Arisemus WI-B09
              |--*A. boxi WM96
              |--A. amydrus WI-B09
              |--A. buzbyae Wagner & Masteller 1996 WM96
              |--A. mariannae Wagner & Masteller 1996 WM96
              `--A. waidei Wagner & Masteller 1996 WM96

Psychodinae incertae sedis:
  Eurygarka helicis QV81
  Breviscapus QV81
  Neotelmatoscopus O98
  Paratelmatoscopus O98
  Epacretron O98
  Gerobrunettia O98
  Notiocharis O98
  Peripsychoda O98
  Ulomyia RD77
  Termitomastus RD77
  Termitodipteron RD77
  Termitadelphos RD77
  Thornburghiella marginalis BW09 [=Pericoma marginalis QV81]

*Type species of generic name indicated

REFERENCES

[BW09] Buck, M., N. E. Woodley, A. Borkent, D. M. Wood, T. Pape, J. R. Vockeroth, V. Michelsen & S. A. Marshall. 2009. Key to Diptera families—adults. 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. 1 pp. 95–156. NRC Research Press: Ottawa.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

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

[M83] Martin, N. A. 1983. Miscellaneous observations on a pasture fauna: an annotated species list. DSIR Entomology Division Report 3: 1–98.

[O98] Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill: Leiden.

[QV81] Quate, L. W., & J. R. Vockeroth. 1981. Psychodidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 293–300. Research Branch, Agriculture Canada.

[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.

[WI-B09] Wagner, R., & S. Ibáñez-Bernal. 2009. Psychodidae (sand flies, and moth flies or owl 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. 1 pp. 319–335. NRC Research Press: Ottawa.

[WM96] Wagner, R., & E. C. Masteller. 1996. New moth flies (Diptera: Psychodidae) and a key to species from Puerto Rico. Proceedings of the Entomological Society of Washington 98 (3): 450–464.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Psychodomorpha

Larva of Philorus sp., from the California Department of Fish and Wildlife.

Belongs within: Neoneura.
Contains: Psychodoidea.

The Psychodomorpha is a clade of flies including the moth flies of the Psychodidae and their close relatives. In the circumscription given below, it is supported by molecular data (Wiegmann et al. 2011). Members include the Blephariceridae, a cosmopolitan group of slender, long-legged flies with wings bearing a secondary network of vein-like markings (Colless & McAlpine 1991).

<==Psychodomorpha WT11
    |--+--Kuperwoodidae GE05
    |  |--Hennigmatidae GE05
    |  |--Grauvogelia [Grauvogeliidae] GE05
    |  |    `--G. arzvilleriana GE05
    |  `--+--Psychodoidea WT11
    |     `--Nadiptera [Nadipteridae] GE05
    |          `--N. kaluginae KK02
    `--Blephariceridae [Blepharicerimorpha, Blepharoceridae] WT11
         |--Edwardsina [Edwardsininae] C09
         |    |--E. ferrugiana CM91
         |    `--E. gigantea WT11
         |--Apistomyiinae [Apistomyiini] CM70
         |    |--Apistomyia tonnoiri CM91
         |    |--Parapistomyia CM91
         |    |--Austrocurupira CM91
         |    `--Neocurupira CM70
         `--Blepharicerinae C09
              |  i. s.: Aposonalco amoyote C09
              |         Kelloggina C09
              |         Limonicola lichanos C09
              |--Paltostoma C09 [Paltostomini H81]
              |    |--P. argyrocincta C09
              |    |--P. exserta C09
              |    `--P. schineri C09
              `--Blepharicerini H81
                   |--Bibiocephala grandis H81
                   |--Dioptopsis H81
                   |    |--D. aylmeri H81
                   |    `--D. sequioarum H81
                   |--Blepharicera H81
                   |    |--B. ostensackeni H81
                   |    |--B. tenuipes H81
                   |    `--B. tetrophtalma O98
                   |--Agathon H81
                   |    |--A. comstocki C09
                   |    |--A. doanei H81
                   |    `--A. elegantulus H81
                   `--Philorus H81
                        |--P. californicus H81
                        |--P. jacinto H81
                        `--P. yosemite H81

*Type species of generic name indicated

REFERENCES

[CM70] Colless, D. H., & D. K. McAlpine. 1970. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 656–740. Melbourne University Press.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

[C09] Courtney, G. W. 2009. Blephariceridae (net-winged midges). 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. 1 pp. 237–243. NRC Research Press: Ottawa.

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

[H81] Hogue, C. L. 1981. Blephariceridae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 191–197. Research Branch, Agriculture Canada.

[KK02] Krzemiński, W., & E. Krzemińska. 2002. Rhaetaniidae, a new family of the Diptera from the Upper Triassic of Great Britain (Diptera: Nematocera). Annales Zoologici 52 (2): 211–213.

[O98] Oosterbroek, P. 1998. The Families of Diptera of the Malay Archipelago. Brill: Leiden.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 29 December 2020.

Phlebotominae

Feeding Phlebotomus papatasi, photographed by Frank Collins.

Belongs within: Psychodoidea.

The Phlebotominae, sand flies, are a group of small flies with soil-dwelling larvae and blood-sucking adults. They are characterised by a four-branched Rs vein in the wing, and mouthparts much longer than the height of the head (Quate & Vockeroth 1981).

<==Phlebotominae GE05
    |--Phlebotomites GE05
    |--Libanophlebotomus lutfallahi GE05
    |--Chinius WI-B09
    |--Sergentomyia WI-B09
    |--Brumptomyia mesai WI-B09
    |--Warileya [incl. Hertigia] WI-B09
    |--Edentomyia WI-B09
    |--Lutzomyia QV81
    |    |--L. cruciata WI-B09
    |    |--L. longipalpis WT11
    |    `--L. vexator QV81
    `--Phlebotomus WT11
         |--P. argentipes A71
         |--P. ariasi A71
         |--P. capensis de Meillon 1955 S61
         |--P. duboscqui WT11
         |--P. longipalpis A71
         |--P. major A71
         |--P. orientalis A71
         |--P. papatasi NJ91
         |--P. parfiliewi A71
         |--P. perniciosus A71
         |--P. sergenti A71
         `--P. verrucarum A71

*Type species of generic name indicated

REFERENCES

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

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

[NJ91] Nuttall, P. A., & L. D. Jones. 1991. Non-viraemic tick-borne virus transmission: mechanism and significance. 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. 3–6. SPB Academic Publishing: The Hague.

[QV81] Quate, L. W., & J. R. Vockeroth. 1981. Psychodidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 293–300. Research Branch, Agriculture Canada.

[S61] Stuckenberg, R. R. 1961. Diptera (Nematocera): Thaumaleidae. In: Hanström, B., P. Brinck & G. Rudebeck. South African Animal Life: Results of the Lund University Expedition in 1950–1951 vol. 8 pp. 409–412. Almqvist & Wiksell: Uppsala.

[WI-B09] Wagner, R., & S. Ibáñez-Bernal. 2009. Psychodidae (sand flies, and moth flis or owl 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. 1 pp. 319–335. Ottawa, NRC Research Press.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Psychodoidea

Male Protoplasa fitchii, from Alexander (1981).

Belongs within: Psychodomorpha.
Contains: Phlebotominae, Psychodinae.

The Psychodoidea are a clade uniting the moth flies and sand flies of the Psychodidae with the primitive crane flies of the Tanyderidae. The Psychodidae are mostly small, very hairy flies with broadly ovate wings with veins M and often Rs each bearing four branches (Colless & McAlpine 1991).

Synapomorphies (from Blagoderov et al. 2002): Ocelli absent; sc-r terminal; M₂ convex; male genitalia inverted.

<==Psychodoidea BLM02
    |--Tanyderidae [Tanyderoidea, Tanyderomorpha] BLM02
    |    |--Praemacrochile BLM02
    |    |--Peringueyomyina A81
    |    |--Protoplasa fitchii A81
    |    |--Nothoderus australiensis CM91, M81
    |    |--Protanyderus margarita A81
    |    |--Macrochile A81
    |    |--Radinoderus occidentalis CM91
    |    `--Eutanyderus A81
    |         |--E. oreonympha CM91
    |         `--E. wilsoni CM91
    `--Psychodidae [Phlebotomidae] WI-B09
         |  i. s.: Eopericoma zherichini Kalugina in Kalugina & Kovalev 1985 RJ93
         |         Tanypsycha GE05
         |         Liassopsychodina GE05
         |         Psychotanyderus GE05
         |         Bazarella WI-B09
         |         Quatiella interdicta WM96
         |--Nannotanyderus WT11
         `--+--Phlebotominae WI-B09
            |--Psychodinae WI-B09
            |--Horaiella [Horaiellinae] WI-B09
            |--Sycoracinae WI-B09
            |    |--Sycorax WI-B09
            |    |--Aposycorax chilensis WI-B09
            |    `--Parasycorax WI-B09
            |--Bruchomyiinae WI-B09
            |    |--Bruchomyia WI-B09
            |    `--Nemapalpus WI-B09
            |         |--N. nearcticus QV81
            |         `--N. youngi WI-B09
            `--Trichomyia [Trichomyiinae] WI-B09
                 |--T. botosaneanui WM96
                 |--T. buchholzi Wagner & Masteller 1996 WM96
                 |--T. flinti Wagner & Masteller 1996 WM96
                 |--T. masneri WM96
                 |--T. nuda WI-B09
                 |--T. rawlinsi WM96
                 |--T. sattelmairi Wagner & Masteller 1996 WM96
                 |--T. urbica QV81
                 `--T. wirthi WM96

*Type species of generic name indicated

REFERENCES

[A81] Alexander, C. P. 1981. Tanyderidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 149–151. Research Branch, Agriculture Canada.

[BLM02] Blagoderov, V. A., E. D. Lukashevich & M. B. Mostovski. 2002. Order Diptera Linné, 1758. The true flies (=Muscida Laicharting, 1781). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 227–240. Kluwer Academic Publishers: Dordrecht.

[CM91] Colless, D. H., & D. K. McAlpine. 1991. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2^nd ed. vol. 2 pp. 717–786. Melbourne University Press: Carlton (Victoria).

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

[M81] McAlpine, J. F. 1981. Morphology and terminology—adults. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 9–63. Research Branch, Agriculture Canada.

[QV81] Quate, L. W., & J. R. Vockeroth. 1981. Psychodidae. In: McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J. Teskey, J. R. Vockeroth & D. S. Wood (eds) Manual of Nearctic Diptera vol. 1 pp. 293–300. Research Branch, Agriculture Canada.

[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.

[WI-B09] Wagner, R., & S. Ibáñez-Bernal. 2009. Psychodidae (sand flies, and moth flies or owl 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. 1 pp. 319–335. NRC Research Press: Ottawa.

[WM96] Wagner, R., & E. C. Masteller. 1996. New moth flies (Diptera: Psychodidae) and a key to species from Puerto Rico. Proceedings of the Entomological Society of Washington 98 (3): 450–464.

[WT11] Wiegmann, B. M., M. D. Trautwein, I. S. Winkler, N. B. Barr, J.-W. Kim, C. Lambkin, M. A. Bertone, B. K. Cassel, K. M. Bayless, A. M. Heimberg, B. M. Wheeler, K. J. Peterson, T. Pape, B. J. Sinclair, J. H. Skevington, V. Blagoderov, J. Caravas, S. N. Kutty, U. Schmidt-Ott, G. E. Kampmeier, F. C. Thompson, D. A. Grimaldi, A. T. Beckenbach, G. W. Courtney, M. Friedrich, R. Meier & D. K. Yeates. 2011. Episodic radiations in the fly tree of life. Proceedings of the National Academy of Sciences of the USA 108 (14): 5690–5695.

Last updated: 29 December 2020.