Tabanomorpha

Los Angeles wormlion Vermileo, photographed by Tokuo Nakamoto.

Belongs within: Orthorrhapha.
Contains: Rhagionidae, Tabanidae.

The Tabanomorpha is a clade of flies united by the presence of a small lobe on the basal segment of the female cercus, and larvae capable of retracting their head into the thorax (Grimaldi & Engel 2005). Many members of this clade are predatory. A number of subgroups, including the Tabanidae, Athericidae and some members of the Spaniidae (previously classified with the Rhagionidae), feed on vertebrate blood (Grimaldi & Engel 2005). Other tabanomorphs include the wormlions of the genus Vermileo whose larvae construct pits like those of ant-lions that they use to capture prey. The Pelecorhynchidae are a group of moderate-sized to large flies whose predatory larvae are found in saturated soils in swamps and stream banks (Teskey 1981a). The Athericidae are medium-sized flies, often with patterned wings, with aquatic larvae mostly inhabiting rocky or gravelly areas of moving streams (Woodley 2009b). The possible stem-athericid Qiyia jurassica is known from larvae from the Middle Jurassic Daohugou Formation of Inner Mongolia that possessed adaptations for ectoparasitism such as piercing mouthparts and a ventral sucker near the front of the thorax (Chen et al. 2014).

Tabanomorpha
    |--+--Rhagionidae WT11
    |  `--Vermileonidae [Protobrachycerontidae, Vermileonoidea] WT11
    |       |--Lampromyia K09
    |       |--Protobrachyceron liasinum RJ93
    |       |--Vermitigris T81b
    |       |    |--V. fairchildi O98
    |       |    |--V. infasciatus O98
    |       |    `--V. orientalis O98
    |       `--Vermileo W09a
    |            |--V. comstocki W09a
    |            |--V. fascipennis W09a
    |            |--V. opacus T81b
    |            |--V. tibialis T81b
    |            `--V. willetti W09a
    `--+--+--Bolbomyia WT11
       |  |    |--B. macgillisi JT81
       |  |    `--B. nana WT11
       |  `--Spaniidae [Spaniinae] GE05
       |       |--Ptiolina majuscula K09, JT81
       |       |--Litoleptis alaskensis K09, JT81
       |       |--Spania nigra K09, JT81
       |       |    |--S. n. nigra JT81
       |       |    `--S. n. americana JT81
       |       |--Symphoromyia GE05
       |       |    |--S. hirta JT81
       |       |    |--S. (Axinicera) inurbana JT81
       |       |    `--S. montana M81
       |       `--Chrysopilus WT11
       |            |--C. ferruginosus O98
       |            |--C. flavibarbis K09
       |            |--C. gratiosus CM91
       |            |--C. ornatus K09
       |            |--C. proximus K09
       |            `--C. thoracicus WT11
       `--+--Batgana GE05
          |--Shulmas GE05
          `--+--Pelecorhynchidae WT11
             |    |--Bequaertomyia jonesi T81a
             |    |--Pseudoerinna K09
             |    |--Pelecorhynchus Macq. 1850 [=Caenopnyga Thoms. 1868] R00
             |    |    |--*P. maculipennis Macq. 1850 [=*Caenopnyga maculipennis] R00
             |    |    |--P. aurantiacus Ricardo 1900 R00
             |    |    |--P. darwini Ricardo 1900 R00
             |    |    |--P. eristaloides (Walker 1848) [=Silvius eristaloides] R00
             |    |    |--P. fulvus CM91
             |    |    |--P. fusconiger (Walker 1848) [=Silvius fusconiger] R00
             |    |    `--P. personatus (Walker 1848) [=Silvius personatus; incl. P. ornatus Schiner 1866] R00
             |    `--Glutops K09
             |         |--G. bandus T81a
             |         |--G. rossi T81a
             |         `--G. singularis WT11
             `--+--Oreoleptis [Oreoleptidae] WT11
                |    `--O. torrenticola WT11
                `--+--Tabanidae WT11
                   `--Athericidae WT11
                        |  i. s.: Atrichops O98
                        |--Qiyia Chen, Wang et al. 2014 CW14
                        |    `--*Q. jurassica Chen, Wang et al. 2014 CW14
                        `--+--Dasyomma [Dasyommatinae] W09b
                           `--Athericinae W09b
                                |--Xeritha W09b
                                |--Suragina W09b
                                |    |--S. concinna W09b
                                |    |--S. latipennis W09b
                                |    `--S. longipes W09b
                                `--Atherix W09b
                                     |--A. ibis O98
                                     |    |--A. i. ibis O98
                                     |    `--A. i. japonica O98
                                     |--A. lantha W81
                                     |--A. maculata [=Rhagio (Atherix) maculatus] G20
                                     |--A. peringueyi S61
                                     `--A. variegata W09b

*Type species of generic name indicated

REFERENCES

[CW14] Chen, J., B. Wang, M. S. Engel, T. Wappler, E. A. Jarzembowski, H. Zhang, X. Wang, X. Zheng & J. Rust. 2014. Extreme adaptations for aquatic ectoparasitism in a Jurassic fly larva. eLife 3: e02844.

[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.

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

[JT81] James, M. T., & W. J. Turner. 1981. Rhagionidae. 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. 483–488. Research Branch, Agriculture Canada.

[K09] Kerr, P. H. 2009. Rhagionidae (snipe 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. 485–489. NRC Research Press: Ottawa.

[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.

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

[R00] Ricardo, G. 1900. Notes on the Pangoninae of the family Tabanidae in the British Museum collection. Annals and Magazine of Natural History, series 7, 5: 97–121, pl. 1.

[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.

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

[T81a] Teskey, H. J. 1981a. Pelecorhynchidae. 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. 459–461. Research Branch, Agriculture Canada.

[T81b] Teskey, H. J. 1981b. Vermileonidae. 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. 529–532. Research Branch, Agriculture Canada.

[W81] Webb, D. W. 1981. Athericidae. 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. 479–482. Research Branch, Agriculture Canada.

[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.

[W09a] Woodley, N. E. 2009a. Vermileonidae (worm-lions). 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. 481–483. NRC Research Press: Ottawa.

[W09b] Woodley, N. E. 2009b. Athericidae (athericid 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. 491–493. NRC Research Press: Ottawa.

Last updated: 12 April 2022.