ISSN 1911-2173

A product of the Biological Survey of Canada & the Entomological Society of Canada

An illustrated identification key to the genera of Ulidiidae (Diptera: Tephritoidea) of the United States and Canada

An illustrated identification key to the genera of Ulidiidae (Diptera: Tephritoidea) of the United States and Canada

Charles Wallace

Biology, School of Life Sciences, The College of Liberal Arts & Science,
Arizona State University, Tempe, Arizona, 85281, United States. [email protected].


Ulidiidae, the "picture-winged flies,'' are a family of true flies belonging to the superfamily Tephritoidea. Over 130 of the family's approximately 700 species, and 40 of its 128 genera, have been documented north of Mexico in the United States and Canada. Although the family is primarily saprophagous and commonly found in association with feces or decaying plant tissue, several species are known to be pestiferous on crops significant to agribusiness, primarily in the American southeast. Here the taxonomic key provided in the "Otitidae" chapter of the Manual of Nearctic Diptera (Steyskal 1987) is updated with color photographs, including notes on taxonomy and distribution of species. Specimens from collections at North Carolina State University, the Florida State Collection of Arthropods, and the National Museum of Natural history were photographed using a Canon DSLR and macro lens, and focus stacked using Zerene Stacker. Information on distribution of species was taken from collection records, online databases, and previous publications.

Steyskal, G.C. 1987. Otitidae. In Manual of Nearctic Diptera. Volume 2. Edited by J.F. McAlpine. Agriculture Canada, Hull, Quebec, Canada. Pp: 799-808.


Tritoxa cuneata species group (photo by C. Wallace)


The "picture-winged flies" (Ulidiidae) are one of the larger families of the Diptera superfamily Tephritoidea, numbering approximately 800 species total, about 135 of which are found in the United States or Canada (Steyskal 1987, Diaz-Fleischer et al. 2000) (some uncertainty remains regarding exact number, due to limited research, taxonomic revisions, and high number of Euxesta Loew). They are a family of acalyptrate flies, generally small to medium, and broadly recognizable by the distinctively patterned wings that grant the family its common name, though this trait is neither universal among nor exclusive to them. Traits common to the family are illustrated in Figure 1.

Figure 1. Lateral habitus of Xanthacrona bipustulata from the Smithsonian National Museum of Natural History collection, showing morphological traits common to Ulidiidae.

Currently, Ulidiidae is divided into two subfamilies: Ulidiinae and Otitinae, each with three tribes (and an additional incertae sedis group of genera in the latter) (Kameneva and Korneyev 2006). These subfamilies are unequally distributed geographically; Otitinae is primarily Holarctic, and represents the slight majority of species documented north of Mexico, whereas Ulidiinae is primarily concentrated in the Neotropical region of Central and South America (Arnett 2000, Kameneva and Korneyev 2006, Galinskaya et al. 2014).

Unlike its close relative, Tephritidae, the ulidiids are primarily a saprophagous group (Ferrar 1987; Arnett 2000). The greater number of picture winged flies can be found in association with feces or, more typically, rotting vegetables and fruits, where they commonly oviposit. Some publications (e.g. Arnett 2000) present the family as exclusively associated with plants, but, although there is a clear preference throughout the family for plant tissue, dead or alive, it is not exclusive (Ferrar 1987, Sivinski 2000, Marshall 2012). There is, however, a repeated trend within the Ulidiidae of true and/or opportunistic phytophagy, in which larvae act as primary, secondary, or tertiary invaders on living plant tissue (Kameneva and Korneyev 2010; Goyal et al. 2012). This is particularly well documented in those species that act as pests on plants such as sugar beet, onions, or corn (Ferrar 1987; Bjerke et al. 1992; Goyal et al. 2010, 2011, 2012).

Like the Tephritidae, adult ulidiids commonly use their distinctive wings to engage in often complex mating rituals. Male members of Callopistromyia Hendel raise their dappled wings at a 90 degree angle over their abdomen and "strut," while others such as Delphinia picta Fabricius make a "rowing" motion while walking over leaves (Sivinski 2000, Marshall 2012). Even more "extreme" examples of novel reproductive strategies can be found in species outside of a Nearctic distribution, as in the stalk-eyed Plagiocephalus Wiedemann, or the post-copulatory ejaculate expulsion in females of Euxesta bilimeki Hendel (Brunel and Rull 2010, Luis Rodriguez-Enriquez et al. 2013).

Unfortunately, the biology of many ulidiids, even in the relatively well-documented North American taxa, remains unknown or understudied. One potential barrier is the near absence of taxonomic tools. There is so far no comprehensive illustrated guide to the taxa of ulidiids of any biogeographical region, and many described species and genera remain to be photographed or otherwise imaged, or else are unavailable to a broad audience. This update to the key will fill some of this void by providing new images of many more taxa than have previously been available.

Materials & Methods

Specimens and data records were accessed from insect collections at three institutions: the Insect Museum in the Department of Entomology and Plant Pathology at North Carolina State University (NCSU), the Florida State Collection of Arthropods (FSCA) at the University of Florida at Gainesville, and the Diptera Collection in the U.S. National Entomological Collection (USNM) at the National Museum of Natural History (NMNH) of the Smithsonian Institution in Washington, DC.

Location data for taxa represented in the United States and Canada was based primarily on Steyskal's (1987) descriptions and supplemented with documentation of occurrences in published literature (Arnett 2000, Marshall 2012), the Catalog of Life (Roskov et al. 2017), the website Systema Dipterorum (Pape and Thompson 2017), the Global Biodiversity Information Facility (GBIF), and collection information from the three institutions listed above. Based on available information, none of the represented species are definitively known from Alaska.

In total, Ulidiidae is represented in the United States and Canada by 40 genera and 132 species. Members of all recognized subfamilial, suprageneric categories (Otitinae: Cephaliini, Myennidini, and Otitini; Ulidiinae: Lipsanini, Pterocallini, Seiopterini, Ulidiini), as of the most recent comprehensive taxonomic consideration of the family (Kameneva and Korneyev 2006), are represented by at least one genus, with a bias towards the primarily Holarctic subfamily Otitinae. A list of genera and species with a documentation in North America north of Mexico is available in Table 1, which also specifies the source (i.e. NCSU) of the specimens photographed for this paper.

Additional Notes

In Arnett (2000), Pseudotephritina Malloch is treated as a sub-genus of Pseudotephritis Johnson. Herein, however, I agree with Steyskal (1987) and Kameneva and Korneyev (2006), in treating it as a genus.

One additional genus, Prionella Robineau-Desvoidy, is indicated as appearing in the United States in Arnett (2000), and on Systema Dipterorum ( However, it is a genus of unrecognizable identity and is therefore excluded.

The terminology used follows the standard set in the "Adult Morphology and Terminology" chapter of the Manual of Afrotropical Diptera (Cumming and Wood 2017). "Frontal index" refers to the ratio of frons height to frons width at base. Figures 2-7 provide pictures of ulidiids from the key with relevant morphological traits labeled.

The bulk of the text in the original dichotomous key is preserved from Steyskal (1987). Where relevant, terms are updated to comply with the terminology from the Manual of Afrotropical Diptera, as in the replacement of "bristle" with "seta." Additional small changes to the original text were made to clarify, correct, or remove redundant and potentially confusing traits. For example, in couplet #30, 30' stated "tibiae usually distinctly banded," but this is contradicted by couplet 32, which clarifies that neither Pseudotephritina Malloch nor Xanthacrona Wulp possess distinctly banded tibiae.

Specimen Imaging

With the exception of Hiatus fulvipes Cresson (represented by non-photographic illustrations) and Texasa chaetifrons Steyskal (type: United States Natural Museum, image: Dr. Allen Norrbom), all images in this key were taken by the author. Each final image was produced through focus stacking. Initial photographs were taken using a Canon EOS 6D camera with either the Canon MP-E 65mm 1-5x macro lens or the Laowa 60mm f/2.8 2X Ultra-Macro (used for some of the specimens from the FSCA), at variable magnification and aperture depending on specimen, mounted on a Cognisys StackShot standard macro rail. Images were stacked using the Zerene Stacker system. Additional images of Texasa chaetifrons, Physiphora clausa Macquart, and Tritoxa flexa Wiedemann were taken with Canon EOS 5DS and MP-E 65mm lens. Edits, to remove blemishes and to correct for color and clarity, were performed using Adobe Photoshop CS6. The specimen identified as Tritoxa cuneata Loew was photographed before the publication by Sinclair et al. (2021) describing two new species in the T. cuneata species complex; locality information and male genitalia are unavailable to determine whether it is actually T. californica Sinclair et al. or T. decipiens Sinclair et al. rather than T. cuneata. One species, Hiatus fulvipes, is known exclusively by several type specimens and was unavailable for photography; it is represented with drawn illustrations. These illustrations were completed by an artist using Plate VI of Cresson (1906) as a reference.

For each trait used diagnostically in the key, an image was attempted; however, several traits were not visible from the available specimens due to the position in which they were pinned.


Table 1. List of all contemporary species of Ulidiidae with a Nearctic distribution. Additional columns indicate species presence in Canada or the United States, as well as origin of specimen photographed ("---" indicates absence of photograph). Abbreviations for repositories: NCSU - Insect Museum in the Department of Entomology and Plant Pathology at North Carolina State University; FSCA - the Florida State Collection of Arthropods at the University of Florida at Gainesville; USNM - Diptera Collection in the U.S. National Entomological Collection at the National Museum of Natural History of the Smithsonian Institution in Washington, DC.

Subfamily Tribe Genus Species# Species Canada USA Specimen source USNMENT #
Otitinae Cephaliini Cephalia 2          
        Cephalia flavoscutellata Becker, 1900 Yes No USNM 1396546
        Cephalia rufipes Meigen, 1826 No Yes USNM 1396541
    Delphinia 1          
        Delphinia picta Fabricius, 1781 Yes Yes NCSU  
    Myiomyrmica 1          
        Myiomyrmica fenestrata Coquillett, 1900 No Yes FSCA, USNM 1396545
    Myrmecothea 1          
        Myrmecothea myrmecoides Loew, 1860 No Yes NCSU, USNM 1396639
    Tritoxa 7          
        Tritoxa californica Sinclair et al. 2021 No Yes ---  
        Tritoxa cuneata Loew, 1873 Yes Yes FSCA  
        Tritoxa decipiens Sinclair et al. 2021 Yes Yes ---  
        Tritoxa flexa Wiedemann, 1830 Yes Yes FSCA  
        Tritoxa incurva Loew, 1873 No Yes FSCA  
        Tritoxa pollinosa Cole, 1919 No Yes FSCA  
        Tritoxa ra Harriot, 1942 No Yes ---  
  incertae sedis Curranops 2          
        Curranops apicalis Cole & Lovett, 1921 Yes Yes ---  
        Curranops scutellaris Coquillett, 1900 No Yes USNM  
    Diacrita 2          
        Diacrita costalis Gerstaecker, 1860 No Yes FSCA, USNM 1396572
        Diacrita plana Steyskal, 1947 No Yes USNM 1396516
    Haigia 1          
        Haigia nevadana Steyskal, 1961 No Yes USNM 1396491
    Idana 1          
        Idana marginata Say, 1830 Yes Yes FSCA  
    Psaeropterella 2          
        Psaeropterella macrocephala Hendel, 1914 No Yes USNM 1396496
        Psaeropterella punctifrons Hendel, 1914 Yes No USNM 1396536
    Tetropismenus 1          
        Tetropismenus hirtus Loew, 1876 No Yes USNM 1396538
    Tujunga 1          
        Tujunga mackenziei Steyskal, 1961 No Yes USNM 1396523
  Myennidini Callopistromyia 2          
        Callopistromyia annulipes Macquart, 1855 No Yes FSCA  
        Callopistromyia strigula Loew, 1873 Yes Yes USNM 1396577
    Dyscrasis 1          
        Dyscrasis hendeli Aldrich, 1932 Yes Yes USNM 1396493
    Oedopa 2          
        Oedopa ascriptiva Hendel, 1909 No Yes USNM 1396533
        Oedopa capito Loew, 1868 No Yes USNM 1396612
    Paroedopa 1          
        Paroedopa punctigera Coquillett, 1900 No Yes USNM 1396531
    Pseudotephritina 2          
        Pseudotephritina cribellum Loew, 1873 Yes Yes FSCA, USNM 1396597
        Pseudotephritina inaequalis Malloch, 1931 No Yes FSCA, USNM 1396629
    Pseudotephritis 4          
        Pseudotephritis approximata Banks, 1914 No Yes FSCA  
        Pseudotephritis corticalis Loew, 1873 Yes Yes FSCA  
        Pseudotephritis inaequalis Malloch, 1931 No Yes ---  
        Pseudotephritis vau Say, 1830 Yes Yes USNM 1396607
    Stictomyia 2          
        Stictomyia longicornis Bigot, 1885 No Yes USNM 1396474
        Stictomyia punctata Coquillett, 1900 No Yes USNM 1396473
    Texasa 1          
        Texasa chaetifrons Steyskal, 1961 No Yes USNM  
    Ulidiotites 1          
        Ulidiotites dakotana Steyskal, 1961 No Yes USNM 1396488
  Otitini Ceroxys 1          
        Ceroxys latiusculus Loew, 1873 Yes Yes FSCA  
    Herina 4          
        Herina canadensis Johnson, 1902 Yes Yes USNM 1396600
        Herina narytia Walker, 1849 No Yes USNM 1396562
        Herina nigribasis McAlpine, 1951 Yes Yes USNM 1396602
        Herina ruficeps Wulp, 1867 Yes Yes FSCA  
    Hiatus 1          
        Hiatus fulvipes Cresson, 1906 No Yes ---  
    Melieria 6          
        Melieria cana Loew, 1858 Yes Yes FSCA  
        Melieria occidentalis Coquillett, 1904 Yes Yes FSCA  
        Melieria ochricornis Loew, 1873 Yes Yes FSCA  
        Melieria picta Meigen, 1826 No Yes USNM 1396486
        Melieria sabuleti Steyskal, 1962 No Yes USNM 1396501
        Melieria similis Loew, 1873 Yes Yes USNM 1396632
    Otites 7          
        Otites bimaculata Hendel, 1911 Yes Yes USNM 1396585
        Otites erythrocephala Hendel, 1911 No Yes USNM 1396590
        Otites erythrosceles Steyskal, 1966 No Yes ---  
        Otites michiganus Steyskal, 1966 Yes Yes FSCA  
        Otites pyrrhocephala Loew, 1876 No Yes USNM 1396634
        Otites snowi Cresson, 1924 No Yes USNM 1396617
        Otites stigma Hendel, 1911 Yes Yes FSCA  
    Tetanops 7          
        Tetanops cazieri Harriot, 1942 No Yes USNM 1396528
        Tetanops integer Loew, 1873 Yes Yes USNM 1396625
        Tetanops luridipennis Loew, 1873 No Yes FSCA  
        Tetanops magdalenae Cresson, 1924 No Yes USNM 1396624
        Tetanops myopaeformis Roder, 1881 Yes Yes FSCA  
        Tetanops parallelus Steyskal, 1970 No Yes USNM 1396587
        Tetanops vittifrons Wulp, 1899 No Yes ---  
Ulidiinae Lipsanini              
    Acrosticta 6          
        Acrosticta apicalis Williston, 1896 No Yes USNM 1396570
        Acrosticta compta Cole, 1912 No Yes ---  
        Acrosticta dichroa Loew, 1874 No Yes USNM 1396503
        Acrosticta fulvipes Coquillett, 1900 No Yes USNM 1396511
        Acrosticta rubida Loew, 1876 No Yes USNM 1396535
        Acrosticta rufiventris Hendel, 1910 No Yes USNM 1396475
    Axiologina 1          
        Axiologina ferrumequinum Hendel, 1909 No Yes FSCA  
    Chaetopsis 7          
        Chaetopsis aenea Wiedemann, 1830 No Yes FSCA  
        Chaetopsis apicalis Johnson, 1900 No Yes FSCA  
        Chaetopsis duplicata Johnson, 1921 No Yes ---  
        Chaetopsis fulvifrons Macquart, 1855 Yes Yes FSCA  
        Chaetopsis magna Cresson, 1924 No Yes ---  
        Chaetopsis massyla Walker, 1849 Yes Yes FSCA  
        Chaetopsis quadrifasciata Curran 1928 No Yes USNM 1396490
    Eumetopiella 2          
        Eumetopiella rufipes Macquart, 1847 No Yes FSCA  
        Eumetopiella varipes Loew, 1866 No Yes USNM 1396506
    Euxesta 33          
        Euxesta abana Curran, 1935 No Yes ---  
        Euxesta abdominalis Loew, 1868 No Yes FSCA  
        Euxesta albitarsis Zetterstedt, 1838 Yes Yes ---  
        Euxesta anna Harriot, 1942 No Yes ---  
        Euxesta annonae Fabricius, 1794 No Yes FSCA  
        Euxesta atlantica Ahlmark , 1995 No Yes USNM 1396576
        Euxesta basalis Walker, 1853 No Yes FSCA  
        Euxesta bicolor Cresson, 1906 No Yes ---  
        Euxesta brookmani Harriot, 1942 No Yes ---  
        Euxesta contorta Curran, 1935 No Yes USNM 1396631
        Euxesta eluta Loew, 1868 No Yes FSCA  
        Euxesta fervida Curran, 1935 No Yes USNM 1396566
        Euxesta fulvicornis Bigot, 1886 No Yes USNM 1396601
        Euxesta juncta Coquillett, 1904 No Yes USNM 1396556
        Euxesta luteocesta No Yes USNM 1396561
        Euxesta lutzi Curran, 1935 No Yes USNM 1396551
        Euxesta magdalenae Cresson, 1924 No Yes ---  
        Euxesta minor Cresson, 1906 No Yes USNM 1396621
        Euxesta nigriceps Curran, 1935 No Yes ---  
        Euxesta nitidiventris Loew, 1873 No Yes USNM 1396586
        Euxesta notata Wiedemann, 1830 No Yes FSCA  
        Euxesta pacifica Steyskal , 1995 No Yes USNM 1396581
        Euxesta pechumani Curran, 1938 No Yes FSCA  
        Euxesta pulchella Cresson, 1906 No Yes USNM 1396596
        Euxesta quaternaria Loew, 1868 No Yes FSCA  
        Euxesta rubida Curran, 1935 No Yes USNM 1396591
        Euxesta sanguinea Hendel, 1913 No Yes USNM 1396616
        Euxesta scutellaris Curran, 1935 No Yes USNM 1396611
        Euxesta spoliata Loew, 1868 No Yes USNM 1396606
        Euxesta thomae Loew, 1868 No Yes ---  
        Euxesta willistoni Coquillett, 1900 No Yes ---  
        Euxesta xeres Curran, 1935 No Yes ---  
        Euxesta zacki Steyskal, 1986 No Yes ---  
    Notogramma 2          
        Notogramma cimiciformis Loew, 1868 No Yes FSCA  
        Notogramma purpuratum Cole, 1923 No Yes FSCA  
    Stenomyia 3          
        Stenomyia hendeli Johnson, 1913 No Yes USNM 1396500
        Stenomyia nasoni Cresson 1913 No Yes USNM 1396582
        Stenomyia tenuissima Hendel, 1910 No Yes FSCA  
    Zacompsia 1          
        Zacompsia fulva Coquillett, 1901 No Yes FSCA  
    Xanthacrona 1          
        Xanthacrona bipustulata Wulp, 1899 No Yes USNM 1396615
    Homalocephala 3          
        Homalocephala apicalis Wahlberg, 1839 Yes Yes USNM 1396470
        Homalocephala bipunctata Loew, 1854 Yes Yes ---  
        Homalocephala similis Cresson, 1924 Yes Yes ---  
    Pseudoseioptera 2          
        Pseudoseioptera albipes Cresson, 1919 Yes Yes USNM 1396505
        Pseudoseioptera dubiosa Johnson, 1921 Yes Yes ---  
    Seioptera 3          
        Seioptera colon Loew, 1868 No Yes USNM 1396513
        Seioptera costalis Walker, 1849 Yes No ---  
        Seioptera vibrans Linnaeus, 1758 Yes Yes USNM 1396515
    Physiphora 6          
        Physiphora alceae Preyssler, 1791 Yes Yes FSCA  
        Physiphora clausa Macquart, 1843 Yes Yes FSCA  
        Physiphora hendeli Johnson, 1913 No Yes ---  
        Physiphora laticauda Loew, 1873 No Yes ---  
        Physiphora nasoni Cresson, 1913 No Yes ---  
        Physiphora tenuis Loew, 1868 No Yes ---  
    Steneretma 1          
        Steneretma laticauda Loew, 1873 No Yes USNM  




Figure 2. Physiphora alceae from the Florida State Collection of Arthropods labeled with directional terms used throughout the text. The double-headed arrow indicates directional terms for dorsal views.

Figure 3. Ceroxys latiusculus from the Florida State Collection of Arthropods, labeled with terms for different characters of the head.

Figure 4. Ceroxys latiusculus from the Florida State Collection of Arthropods, labeled with terms for different characters of the head.

Figure 5. Tritoxa flexa from the Florida State Collection of Arthropods, with labels for characters visible dorsally.

Figure 6. Melieria picta from the Smithsonian National Museum of Natural History Collection with thoracic setae used in this key labeled.

Figure 7. Wing of Ceroxys latiusculus from the Florida State Collection of Arthropods with veins, crossveins, and cells used for identification in this key labeled.

Species Keys


I would like to thank Dr. Gary Steck at the Florida State Collection of Arthropods for facilitating access to the FSCA in Gainesville, and to Dr. Torsten Dikow and Dr. Allen Norrbom for facilitating access to the USNM collection at the Smithsonian. Thanks to Dr. Allen Norrbom for photographing Texasa chaetifrons, whose type is held in the USNM collection, and Alyssa Seemann for help with additional photographs of Tritoxa flexa, Physiphora clausa, Tetropismenus hirtus and Texasa chaetifrons in the USNM collection. Additionally, thanks to those involved with the Insect Museum and Insect Systematics program at North Carolina State University: especially, Bob Blinn, the collection manager; Dr. Matthew Bertone; and my MS adviser, Dr. Brian Wiegmann. Also to Kees Eglowski for illustrating Hiatus fulvipes. And thanks to the late Dr. George C. Steyskal, for all his exceptional scholarship and foundational work in fly taxonomy.


Allen, E.J. and Foote, B.A. 1967. Biology and Immature Stages of Three Species of Otitidae (Diptera) Which Have Saprophagous Larvae. Annals of the Entomological Society of America, 60: 826-836.

Arnett, R.H. 2000. American Insects: A Handbook of the Insects of America North of Mexico. CRC Press, Boca Raton, Florida, United States.

Bjerke, J.M., Anderson, A.W. and Freeman, T.P. 1992. Morphology of the Larval Stages of Tetanops myopaeformis (Roder) (Diptera: Otitidae). Journal of the Kansas Entomological Society, 65: 59-65. Available from [Accessed 19 April 2021].

Brunel, O. and Rull, J. 2010. Natural History and Mating Behavior of Pseudodyscrasis scutellaris, a Fly Species (Ulidiidae) Associated With Agave in Mexico. Annals of the Entomological Society of America, 103: 430-438.

Cresson, E.T. 1906. Some North American Diptera from the South West Paper I. Transactions of the American Entomological Society (1890-), 32: 279-288.

Cumming, J.M., and Wood, M.D. 2017. Adult Morphology and Terminology. In Manual of Afrotropical Diptera. Edited by Kirk-Spriggs, A.H., and Sinclair, B.J. South African National Biodiversity Institute, Pretoria, Gauteng, South Africa. Pp: 89-134.

Diaz-Fleischer, F., and Aluja, M. 2000. Behavior of tephritid fies: a historical perspective. In Fruit Fies (Tephritidae): phylogeny and evolution of behavior. Edited by M. Aluja and A. L. Norrbom. CRC, Boca Raton, Florida, United States. Pp. 39-69.

Ferrar, P. 1987. Family Otitidae. In A Guide to the Breeding Habits and Immature Stages of Diptera Cyclorrhapha. Edited by L. Lyneborg. Scandinavian Science Press Ltd, Vinderup, Denmark. Pp 266-270.

Galinskaya, T.V., Suvorov, A., Okun, M.V., and Shatalkin, A.I. 2014. DNA barcoding of Palaearctic Ulidiidae (Diptera: Tephritoidea): Morphology, DNA evolution, and Markov codon models. Zoological Studies, 53: 1-17.

Goyal, G., Nuessly, G.S., Steck, G.J., Seal, D.R., Capinera, J.L., and Boote. K.J. 2010. New Report of Chaetopsis massyla (Diptera: Ulidiidae) as a Primary Pest of Corn in Florida. Florida Entomologist, 93: 198-202.

Goyal, G., Nuessly, G.S., Seal, D.R., Capinera, J.L., Steck, G.J., and Boote, K.J. 2011. Distribution of Picture-Winged Flies (Diptera: Ulidiidae) Infesting Corn in Florida. Florida Entomologist, 94: 35-47.

Goyal, G., Nuessly, G.S., Steck, G.J., Seal, D.R., Capinera, J.L., and Meagher, R.L. 2012. Examination of the Pest Status of Corn-Infesting Ulidiidae (Diptera). Environmental Entomology, 41: 1131-1138.

Harriot, S.C. 1942. A New Genus and a New Species of Otitidae from North America (Diptera). Journal of the New York Entomological Society, 50: 249-250.

Kameneva, E.P. and Korneyev, V.A. 2006. Myennidini, a new tribe of the subfamily Otitinae (Diptera: Ulidiidae), with discussion of the suprageneric classification of the family. Biotaxonomy of Tephritoidea. Israel Journal of Entomology, 35-36: 497-586. Available from
[Accessed 19 April 2021].

Kameneva, E.P. and Korneyev, V.A. 2010. Ulidiidae (Picture-Winged Flies). In Manual of Central American Diptera: Volume 2. Edited by B.V. Brown, A. Borken, J.M. Cumming, D.M. Wood,N.E. Woodley, and M.A. Zumbado. NRC Research Press, Ottawa, Ontario, Canada. Pp: 886-902.

Kameneva, E.L., and Korneyev, V.A. 2015. Key to the species of the genus Homalocephala Zetterstedt, 1838 (Diptera: Ulidiidae). Ukrainska Entomofaunistyka, 6: 47-49. Available from [Accessed 19 April 2021].

Kameneva, E.P., and Korneyev, V.A. 2016. Revision of the Genus Physiphora Fallén 1810 (Diptera: Ulidiidae: Ulidiinae). Zootaxa, 4087: 1-88.

Kameneva, E.P., Korneyev, V.A., and Ramos-Pastrana, Y. 2017. A New Genus, New Species and New Records of Ulidiidae (Diptera: Tephritoidea) From Colombia. Vestnik Zoologii, 51: 125-136.

Korneyev, V.A. 2000. Phylogenetic relationships among the families of the superfamily Tephritoidea. In Fruit Fies (Tephritidae): phylogeny and evolution of behavior. Edited by M. Aluja and A. L. Norrbom. CRC, Boca Raton, Florida, United States. Pp 3-22.

Malloch, J. R. 1931. Flies of the genus Pseudotephritis Johnson (Diptera: Ortalidae). Proceedings of the United States National Museum, 79: 1-6.

Marshall, S.A. 2012. Flies: The Natural History and Diversity of Diptera. Firefly Books Inc, Buffalo, New York, United States.

McAlpine, J.F. A Review of the North American Species of Herina Robineau-Desvoidy (= Tephronota Loew) (Diptera: Otitidae). The Canadian Entomologist, 83: 308-314.

Morgulis, E., Freidberg, A., and Kameneva, E.P. 2013. Two new species of Herina (Diptera: Ulidiidae) from the Mediterranean region, with key to species groups. Zootaxa, 3686: 461-470.

Pape, T., and Thompson, F.C. 2017. Systema Dipterorum, Version 1.5 [online]. Available from [accessed 5 December 2017].

Roskov, Y., Abucay, L., Orrell, T., Nicolson, D., Bailly, N., Kirk, P.M., Bourgoin, T., DeWalt, R.E., Decock, W., De Wever, A.. van Nieukerken, E., Zarucchi, J., and Penev, L. 2017. Species 2000 & ITIS Catalogue of Life [online]. Available from [accessed 28 November 2017].

Sinclair, B. J., Macleod, A. M., and Wheeler, T. A. 2021. Revision of the Nearctic genus Tritoxa Loew (Diptera: Ulidiidae). Zootaxa, 4920: 359-379.

Sivinski, J. 2000. Breeding Habits and Sex in Families Closely Related to the Tephritidae: Opportunities for Comparative Studies of the Evolution of Fruit Fly Behavior. In Fruit Fies (Tephritidae): phylogeny and evolution of behavior. Edited by M. Aluja and A. L. Norrbom. CRC, Boca Raton, Florida, United States. Pp 23-38.

Steyskal, G. C. 1941. The genus Diacrita Gerstaecker (Diptera, Otitidae). Bulletin of the Brooklyn Entomological Society, 41: 149-154. Available from [Accessed 19 April 2021].

Steyskal, G. C. 1962. The genus Curranops Harriot (Diptera: Otitidae). Proceedings of the Entomological Society of Washington, 64: 117-118. Available from [Accessed 19 April 2021].

Steyskal, G. C. 1963. The genus Notogramma Loew (Diptera, Acalyptratae, Otitidae). Proceedings of the Entomological Society of Washington, 65: 195-200. Available from [Accessed 19 April 2021].

Steyskal, G. C. 1966a. A review of the North American species of the genus Otites Latreille, with descriptions of two new species (Diptera: Otitidae). The Great Lakes Entomologist, 1: 79-84. Available from[Accessed 18 May 2020].

Steyskal, G.C. 1966b. A key to species of the genus Xanthacrona Wulp (Diptera, Otitidae). Proceedings of the Entomological Society of Washington, 68: 269.

Steyskal, G. C. 1967. A key to the species of Tritoxa Loew (Diptera: Otitidae). Proceedings of the Entomological Society of Washington, 69: 94. Available from [Accessed 19 April 2021].

Steyskal, G. C. 1970. A new North American species of Tetanops Fallén sensu sticto (Diptera, Otitidae). Journal of the Kansas Entomological Society, 43: 378-380. Available from [Accessed 17 May 2020].

Steyskal, G.C. 1971. A new Central American species of Zacompsia Coquillett, with a key to the described species (Diptera: Otitidae). Proceedings of the Entomological Society of Washington, 73: 247-248. Available from [Accessed 19 April 2021].

Steyskal, G.C. 1973. The Genus Axiologina Hendel (Diptera: Otitidae). The Florida Entomologist, 56: 132-134. Available from [Accessed 9 May 2020].

Steyskal, G. C. 1974. The genus Oedopa Loew (Diptera, Otitidae, Ulidiinae). Proceedings of the Entomological Society of Washington, 76:163-164. Available from [Accessed 19 April 2021].

Steyskal, G. C. 1975. The genus Callopistromyia Hendel (Diptera: Otitidae). Entomological News, 85: 147-148. Available from [Accessed 19 April 2021].

Steyskal, G.C. 1987. Otitidae. In Manual of Nearctic Diptera. Volume 2. Edited by J.F. McAlpine. Agriculture Canada, Hull, Quebec, Canada. Pp: 799-808.

Steyskal, G.C. 1991. Description of new species of Otitidae from Chile (Diptera). Acta Entomologica Chilena, 16: 25-28.

Steyskal, G.C., and Ahlmark, K.M. 1995. Two new species of Euxesta Loew (Diptera; Otitidae). Insecta Mundi, 9: 189-193. Available from [Accessed 19 April 2021].


Wallace, C. 2021. An illustrated identification key to the genera of Ulidiidae (Diptera: Tephritoidea) of the United States and Canada. Canadian Journal of Arthropod Identification 45: 94 pp. doi:10.3752/cjai.2021.45