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.

Culicoidini

Culicoides obsoletus, from here.

Belongs within: Ceratopogonidae.

The Culicoidini are a group of biting midges whose females commonly feed on the blood of vertebrates, with some Culicoides species being vectors of parasites or viruses. Some species can be significant pests in the latter regard, with attacks from Culicoides furens in coastal regions in Central America being bad enough to preclude human habitation in some locations (Borkent et al. 2009). Members of the genus Paradasyhelea have the palps and mouthparts reduced relative to Culicoides, and females lack the teeth on the mandible of the latter genus (Downes & Wirth 1981).

Characters (from Downes & Wirth 1981): Female antenna with 13 flagellomeres. Prescutal pits prominent. Wing with macrotrichia usually abundant; cell r₁ and cell r₂₊₃ both usually well-developed, similar in size; M forking beyond crossvein r-m, medial fork petiolate. Empodia small; claws of both sexes small, equal, and simple.

<==Culicoidini BSG09
    |--Paradasyhelea DW81
    |    |--P. minuta DW81
    |    `--P. olympiae DW81
    `--Culicoides BSG09
         |  i. s.: C. angularis CM91
         |         C. anophelis CM91
         |         C. antennalis CM91
         |         C. arubae BSG09
         |         C. austeni A71
         |         C. barbosai BSG09
         |         C. brevitarsis CM91
         |         C. castillae WB56
         |         C. equatoriensis WB56
         |         C. filarifer BSG09
         |         C. foxi BSG09
         |         C. furens BSG09
         |         C. gabaldoni WB56
         |         C. grahami A71
         |         C. impunctatus A71
         |         C. inornatipennis A71
         |         C. insignis BSG09
         |         C. insinuatus Ortiz & Leon 1955 WB56
         |         C. marmoratus CM74
         |         C. ornatus L91
         |         C. paraensis BSG09
         |         C. phlebotomus BSG09
         |         C. pseudodiabolicus BSG09
         |         C. pulicaris A71
         |         C. pusillus BSG09
         |         C. yoosti GE05
         |         C. debilipalpis group WB56
         |           |--C. dasyophrus Macfie 1940 WB56
         |           |--C. debilipalpis [incl. C. debilipalpis var. glabrior Macfie 1940] WB56
         |           |--C. germanus Macfie 1940 WB56
         |           |--C. hoffmani Fox 1946 WB56
         |           `--C. trilineatus Fox 1946 WB56
         |--C. (Culicoides) yukonensis M90
         |--C. (Avaritia) obsoletus DW81
         |--C. (Beltranmyia) crepuscularis DW81
         |--C. (Diphaomyia) haematopotus DW81
         |--C. (Drymodesmyia) copiosus DW81
         |--C. (Monoculicoides) variipennis M90
         `--C. (Selfia) hieroglyphicus DW81

*Type species of generic name indicated

REFERENCES

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

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). 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. 407–435. NRC Research Press: Ottawa.

[CM74] Colless, D. H., & D. K. McAlpine. 1974. Diptera (flies). In: CSIRO. The Insects of Australia: A textbook for students and research workers. Supplement 1974 pp. 91–96. 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).

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. 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. 393–421. Research Branch, Agriculture Canada.

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

[L91] Liehne, P. F. S. 1991. An Atlas of the Mosquitoes of Western Australia. Health Department of Western Australia.

[M90] McAlpine, J. F. 1990. Insecta: Diptera adults. In: Dindal, D. L. (ed.) Soil Biology Guide pp. 1211–1252. John Wiley & Sones: New York.

[WB56] Wirth, W. W., & F. S. Blanton. 1956. Redescriptions of four species of Neotropical Culicoides of the debilipalpis group (Diptera: Heleidae). Journal of the Washington Academy of Sciences 46 (6): 186–190.

Last updated: 28 December 2020.

Forcipomyia

Male Forcipomyia, copyright Ashley Bradford.

Belongs within: Ceratopogonidae.

Forcipomyia is a diverse genus of hairy biting midges.

Characters (from Downes & Wirth 1981): Female antenna with 13 flagellomeres. Wing with C short or long; cell r₂₊₃ usually short, but if long distinctly narrow; crossvein r-m present; microtrichia minute; macrotrichia moderately abundant, sloping, often scale-like, covering most of wing; fringe complex, not a single row of hairs. Empodia well developed in female; claws strongly curved.

<==Forcipomyia
    |  i. s.: F. acidicola BSG09
    |         F. fusculus BR09
    |         F. townsendi A71
    |         F. utae A71
    |--F. (Forcipomyia) bipunctata DW81
    |--F. (Blantonia) caribbea DW81
    |--F. (Caloforcipomyia) glauca DW81
    |--F. (Euprojoannisia) wirthi DW81
    |--F. (Lasiohelea) M81
    |    |--F. (L.) fairfaxensis M81
    |    `--‘Lasiohelea’ opilionivora CM07
    |--F. (Metaforcipomyia) pluvialis DW81
    |--F. (Microhelea) BSG09
    |    |--F. (M.) fuliginosa DW81
    |    `--F. (M.) ixodoides BSG09
    |--F. (Pterobosca) fusicornis DW81
    |--F. (Synthyridomyia) colemani DW81
    |--F. (Thyridomyia) monilicornis DW81
    |--F. (Trichohelea) eques DW81
    `--F. (Warmkea) aeria DW81

*Type species of generic name indicated

REFERENCES

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

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). 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. 407–435. Ottawa, NRC Research Press.

[BR09] Borkent, A., & G. Rotheray. 2009. Key to Diptera families—larvae. 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. 157–191. Ottawa, NRC Research Press.

[CM07] Cokendolpher, J. C., & P. G. Mitov. 2007. Natural enemies. In: Pinto-da-Rocha, R., G. Machado & G. Giribet (eds) Harvestmen: The Biology of Opiliones pp. 339–373. Harvard University Press: Cambridge (Massachusetts).

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. 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. 393–421. Research Branch, Agriculture Canada.

[M81] McAlpine, J. F. 1981. Key to families—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. 89–124. Research Branch, Agriculture Canada.

Sphaeromiini

Mallochohelea larva, from The University of Iowa.

Belongs within: Ceratopogonidae.

The Sphaeromiini are a group of biting midges whose females have a series of stout, blunt spines on the underside of the fifth tarsomere (Downes & Wirth 1981).

Characters (from Downes & Wirth 1981): Female antenna with 13 flagellomeres. Wing with M forking at or before crossvein r-m, medial fork sessile. Tarsomere 5 of female armed ventrally with stout black blunt spines (batonnets). Female abdomen without internal sclerotized gland rods; sternite 8 of female often with pair of hair tufts.

<==Sphaeromiini DW81
    |--Sphaeromias longipennis DW81
    |--Jenkinshelea magnipennis DW81
    |--Macropeza blantoni DW81
    |--Lanehelea BSG09
    |--Neobezzia BSG09
    |--Johannsenomyia DW81
    |    |--J. argentata DW81
    |    `--J. blantoni BSG09
    |--Nilobezzia DW81
    |    |--N. minor DW81
    |    `--N. schwarzii BSG09
    |--Probezzia DW81
    |    |--P. concinna DW81
    |    `--P. pallida DW81
    `--Mallochohelea DW81
         |--M. albibasis DW81
         |--M. pullata BSG09
         `--M. smithi DW81

*Type species of generic name indicated

REFERENCES

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). 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. 407–435. Ottawa, NRC Research Press.

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. 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. 393–421. Research Branch, Agriculture Canada.

Ceratopogonini

Male Ceratopogon cf. inverecundus, copyright NTNU university Museum.

Belongs within: Ceratopogonidae.

The Ceratopogonini are a group of biting midges whose eyes are usually pubescent and wings often milky in appearance (Downes & Wirth 1981).

Characters (from Downes & Wirth 1981): Eye usually pubescent. Female antenna with 13 flagellomeres. Wing often milky. Cell r₂₊₃ usually small, not or only a little longer than cell r₁, one or both cells sometimes lost; crossvein r-m nearly perpendicular to R₄₊₅; M forking beyond crossvein r-m.

<==Ceratopogonini DW81
    |--Alluaudomyia bella DW81
    |--Isohelea stigmalis DW81
    |--Rhynchohelea monilicornis DW81
    |--Ceratoculicoides DW81
    |--Brachypogon DW81
    |    |--B. paraensis DW81
    |    `--B. stigmalis BSG09
    `--Ceratopogon BSG09
         |--C. bicolor K01
         |--C. culicoidithorax DW81
         |--C. inverecundus BSG09
         |--C. sericatus K01
         `--C. willisi BSG09

*Type species of generic name indicated

REFERENCES

[BSG09] Borkent, A., G. R. Spinelli & W. L. Grogan Jr. 2009. Ceratopogonidae (biting midges, purrujas). 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. 407–435. Ottawa, NRC Research Press.

[DW81] Downes, J. A., & W. W. Wirth. 1981. Ceratopogonidae. 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. 393–421. Research Branch, Agriculture Canada.

[K01] Kertész, K. 1901. Legyek [Dipteren]. 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. 179–201. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

Tanytarsini

Micropsectra junci, copyright J. K. Lindsey.

Belongs within: Chironomidae.

The Tanytarsini are a group of non-biting midges with a long first tarsomere on the fore leg and usually with macrotrichia on the wings (Oliver 1981).

Characters (from Oliver 1981): Wing membrane usually with macrotrichia; calypter always without fringe; crossvein r-m nearly parallel to R₄₊₅, and continuous with it, or if oblique, then wing apex truncated; crossvein m-cu absent. First tarsomere of foreleg longer than fore tibia; hind tibia with two combs composed of basally fused spines. Gonostylus fused to gonocoxite and directed backward.

<==Tanytarsini G09
    |--Corynocera ambigua O81, R86
    |--Lenziella cruscula O81
    |--Zavrelia O81
    |--Stempellina bausei O81, R86 [incl. S. montivaga R86]
    |--Lauterbornia O81
    |--Nandeva SA09
    |--Pontomyia SA09
    |--Skutzia SA09
    |--Constempellina SA09
    |--Neostempellina thienemanni SA09, R86
    |--Stempellinella SA09
    |--Neozavrelia luteola SA09
    |--Parapsectra SA09
    |--Sublettea SA09
    |--Virgatanytarsus arduennensis SA09
    |--Rheotanytarsus O81
    |    |--R. calakmulensis SA09
    |    `--R. rhenanus R86
    |--Paratanytarsus O81
    |    |--P. grimmii SA09
    |    `--P. quadratus SA09
    |--Cladotanytarsus O81
    |    |--C. lewisi P09
    |    |--C. mancus D91
    |    `--C. viridiventris O81
    |--Caladomyia SA09
    |    |--C. pistra SA09
    |    `--C. riotarumensis SA09
    |--Tanytarsus SA09
    |    |--T. barbitarsus PH10
    |    |--T. ejuncidus R86
    |    |--T. guatemalensis Sublette & Sasa 1994 SA09
    |    |--T. heusdensis G09
    |    |--T. pallidocornis G09
    |    |--T. pandus SA09
    |    `--T. vespertinus B87
    `--Micropsectra G09
         |--M. atitlanensis Sublette & Sasa 1994 SA09
         |--M. atrofasciata G09
         |--M. bidentata G09
         |--M. junci G09
         |--M. longicrista G09
         |--M. notescens G09
         |--M. pallidula G09
         |--M. schrankelae G09
         `--M. subvirida G09

*Type species of generic name indicated

REFERENCES

[B87] Boothroyd, I. K. G. 1987. Taxonomic composition and life cycles of Chironomidae (Diptera) in a northern New Zealand stream. Entomologica Scandinavica Supplement 29: 15–22.

[D91] Davids, C. 1991. Water mites: the impact of larvae and adults on their host and prey populations. 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. 1 pp. 497–501. SPB Academic Publishing: The Hague.

[G09] Gerecke, R. 2009. Revisional studies on the European species of the water mite genus Lebertia Neuman, 1880 (Acari: Hydrachnidia: Lebertiidae). Abhandlungen der Senckenberg Gesellschaft für Naturforschung 566: 1–144.

[O81] Oliver, D. R. 1981. Chironomidae. 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. 423–458. Research Branch, Agriculture Canada.

[P09] Pape, T. 2009. Economic importance of Diptera. 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. 65–77. Ottawa, NRC Research Press.

[PH10] Pinder, A. M., S. A. Halse, R. J. Shiel & J. M. McRae. 2010. An arid zone awash with diversity: patterns in the distribution of aquatic invertebrates in the Pilbara region of Western Australia. Records of the Western Australian Museum Supplement 78 (1): 205–246.

[R86] Reiss, F. 1986. Erster Nachtrag zur Chironomidenfauna Bayerns (Diptera, Chironomidae). Spixiana 9 (2): 175–178.

[SA09] Spies, M., T. Andersen, J. H. Epler & C. N. Watson Jr. 2009. Chironomidae (non-biting 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. 437–480. Ottawa, NRC Research Press.

Procladiini

Female Procladius, copyright Tom Murray.

Belongs within: Chironomidae.

The Procladiini are a group of non-biting midges characterised by wings with a long stem to the cubital fork, at least half as long as CuA₂, with the cubital fork being distal to crossvein m-cu.

<==Procladiini SA09
    |--Djalmabatista pulchra SA09
    |--Laurotanypus travassosi Oliveira, Messias & da Silva-Vasconcelos 1992 SA09
    `--Procladius SA09
         |  i. s.: P. freemani O81
         |--P. (Procladius) mozambique SA09
         |--P. (Holotanypus) culiciformis SA09
         `--P. (Psilotanypus) SA09
              |--P. (P.) bellus SA09
              |--P. (P.) etatus Roback 1982 SA09
              `--P. (P.) stroudi Roback 1982 SA09

*Type species of generic name indicated

REFERENCES

[O81] Oliver, D. R. 1981. Chironomidae. 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. 423–458. Research Branch, Agriculture Canada.

[SA09] Spies, M., T. Andersen, J. H. Epler & C. N. Watson Jr. 2009. Chironomidae (non-biting 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. 437–480. Ottawa, NRC Research Press.