BIOTAXONOMY OF Isr. J. Entomol. Vol. 35-36, 2005/6, pp. 497-586

Myennidini, a New Tribe of the Subfamily Otitinae (Diptera: ), with Discussion of the Suprageneric Classification of the Family

Elena Kameneva1 and Valery Korneyev2

ABSTRACT The diagnoses of all six tribes of Ulidiidae are revised. The Myennidini, new tribe (subfamily Otitinae), is defined to include 14 genera: Robineau- Desvoidy, Oedopa Loew, Stictomyia Bigot, Paroedopa Coquillett, Johnson, Hendel, Stictoedopa Brèthes, Pseudotephritina Malloch, Aldrich, Ulidiotites Steyskal, Pseudodyscrasis Hernández, Arborotites Barraclough, Namibotites Barraclough, and Neodyscrasis Kameneva and Korneyev, n. gen. (type species: Pseudodyscrasis steyskali Hernández, 1988). These genera were previously erroneously assigned either to the Ulidiini or to the Pterocallini; the latter tribe is shown here to belong to the subfamily . A phylogenetic analysis is provided for the tribes of Ulidiidae and the genera of Myennidini. Most genera are redescribed, and the distributions of all the species are discussed, including new records. The following synonymy is established: Myennis tricolor Hendel, 1909 = Myennis nebulosa Krivosheina and Krivosheina, 1997, n. syn. Pterotaenia Rondani, 1868 is resurrected from synonymy with Macquart.

1Division of General and Applied Entomology, the I.I. Schmalhausen Institute of Zoology, 01601 Kiev, Ukraine. E-mail: @yandex.ru. 2Laboratory of Biogeography, the I.I. Schmalhausen Institute of Zoology, 01601 Kiev, Ukraine. E-mail: [email protected]. BIOTAXONOMY OF TEPHRITOIDEA

INTRODUCTION The family Ulidiidae belongs to the Higher Tephritoidea, a monophyletic group also containing the , and (Korneyev, 1999). Relationships among the higher taxa in this group were discussed by Hennig (1958), J.F. McAlpine (1977, 1989) and, recently, Korneyev (1999), who also summarized our recent studies of suprageneric taxa within the Ulidiidae. Loew (1868a, 1868b) proposed the first detailed classification of the family (as Ortalidae). Within the Ortalidae he recognized seven family-group taxa: Pyrgotina, Ortalina, Cephalina, Platystomina, Pterocallina, Ulidiina and Richardiina. The groups derived from Platystoma Meigen (Platystomatidae, Platystomatinae), Pyrgota Wiedemann (Pyrgotidae, Pyrgotinae, Pyrgotini) and Richardia Robineau-Desvoidy (, Richardiinae) were later accorded family status. The family-group name Ortalidae (Ortalinae, Ortalini) based on Fallén, 1810, a junior homonym of Ortalis Merrem, 1786 (Aves), is unavailable. The other three names are valid names of the family-group taxa in the family Ulidiidae (= Pterocallidae, = Otitidae). Later, the Ulidiidae, Pterocallidae and Ortalidae were often considered as subfamilies of the “Muscaridae” or as separate families (Hendel, 1909a, 1909b, 1909c, 1910b, 1911, 1916). Since the Platystomatidae (as Platystomidae, including Cephaliini), Pyrgotidae, Pterocallidae and Richardiidae were excluded from the family, a new family name, Otitidae, was proposed to replace Ortalidae (Aldrich, 1932), when the name Ortalidae was found to be unavailable. This nomenclatural change was accepted by Malloch (1932, 1940), Curran (1934b, 1938), and Hendel (1936b) and was also used by Hennig (1939, 1940), who treated the Otitidae (including the former Pterocallidae) and Ulidiidae (including the newly established subfamily Euxestinae) as separate families. Thus, some authors, mainly American and Australian (Crampton, 1944; Steyskal, 1961, 1965, 1968, 1987; Colless and D.K. McAlpine, 1970, 1991; J.F. McAlpine, 1977, 1989; Cole, 1969), have traditionally recognized one family name, Otitidae, whereas most European dipterists (Richter, 1970a, 1970b; Soós, 1984; Zaitzev, 1984; Greve, 1998) used two separate family names, Ulidiidae and Otitidae, following Hennig’s earlier opinion. Kameneva and Korneyev (1994) emphasized that the senior valid name for the family that unites both Ulidiidae and Otitidae sensu Hennig (1939, 1940) is Ulidiidae Macquart, 1835 rather than Otitidae Aldrich, 1932. Some dipterists working with the family (Merz, 1996, 1998; Clements and Merz, 1997, 1998; Clements, 1998; Barraclough, 2000) accepted this point of view, while others (D.K. McAlpine, F.C. Thompson, personal communication) insist that conservation of the name Otitidae versus Ulidiidae by the plenary power of the International Commission of Zoological Nomenclature is preferable. The main objection to the latter point of view is that Ulidiidae is certainly not an unused name; and, moreover, it has been in use for more than 50 years along with Otitidae. When the priority of the name Ulidiidae over Otitidae was demonstrated (Kameneva and Korneyev, 1994), application of the former name met the requirements of Article 23 of the third edition of the International Code of Zoological Nomenclature (International Commission on Zoological Nomenclature, 1985), and the principle of priority was applied explicitly to stabilize the situation caused by the prolonged coexistence of two competing names. In our opinion, further changes to the family name would destabilize nomenclature, and contradict Article 23.2 of the fourth edition of the Code (International Commission on Zoological Nomenclature, 1999). The name Tetanopina Loew, 1862, based upon Fallén (closely related to

498 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Latreille), is another name that has priority over Otitidae Aldrich, 1932 (Otitinae, Otitini). However, it has never been used since it was proposed and therefore cannot be accepted as a valid family-group name. Furthermore, the family-group names Cephalinae [sic] Schiner, 1864 and Pterocallina Loew, 1868 also have priority over Otitidae and derived names, although none of them has ever been applied to the family as a whole or to one of the two subfamilies. The suprageneric classification of the Ulidiidae is not well established. Steyskal (1961, 1965, 1982, 1987) divided the family (as Otitidae) into two subfamilies, Ulidiinae and Otitinae, but he did not further subdivide them into tribes. Kameneva and Korneyev (1994) proposed a tribal classification and provided a preliminary analysis of phylogenetic relationships among the suprageneric taxa in Ulidiidae. However, certain statements made in that paper were erroneous. First of all, they considered the Myennis Robineau-Desvoidy as a member of the tribe Pterocallini, and included the latter tribe in the subfamily Otitinae. Our recent studies of the genera assigned to Pterocallini have shown most of the Neotropical representatives of that tribe to be closely related to the genus Rondani, 1848, sharing several genitalic characters with the predominantly Neotropical groups of genera related to Loew, 1868. At the same time, Myennis, Pseudotephritis Johnson, 1902 and related genera were found to possess characters that indicate their relationship to Otitini rather than to Pterocallini. Based on our preliminary results, we stated that the Ulidiidae should be subdivided into two subfamilies: Ulidiinae (with tribes Seiopterini, Ulidiini, Lipsanini (= Euxestini), and Pterocallini), and Otitinae (with tribes Cephaliini, Otitini, and an unnamed tribe that corresponds to the group of genera related to Myennis) (Kameneva and Korneyev in Korneyev, 1999). In the current paper, the latter group of genera is described as a new tribe, Myennidini, and its relationships with the other tribes of the family are discussed.

MATERIALS AND METHODS The examined material of myennidine species is listed in Appendix 1. Detailed label data of other taxa are not included, because they are both irrelevant and too numerous. The following abbreviations refer to institutions where the examined specimens are deposited: AMNH — American Museum of Natural History, New York, USA ANSP — Academy of Natural Sciences of Philadelphia, Pennsylvania, USA CMP — Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, USA CNC — Canadian National Collections of , Arachnids and Nematodes, Ottawa, Canada INBio — INBio, Santo Domingo, Heredia, Costa Rica MCZ — Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA MSNT — Museo Regionale Scienze Naturale, Torino, Italy NHMW — Natural History Museum of Vienna (Naturhistorische Museum Wien), Austria SIZK — Schmalhausen Institute of Zoology, National Academy of Sciences of Ukraine, Kiev, Ukraine TAUI — Tel Aviv University, Israel HNHM — Hungarian Natural History Museum (Természettudományi Muzeum), Budapest, Hungary

499 BIOTAXONOMY OF TEPHRITOIDEA

USNM — National Museum of Natural History, Washington, D.C., USA ZMHB — Museum of Natural History of the Humboldt University in Berlin (Museum für Naturkunde der Humboldt-Universität zu Berlin), Germany ZMUM — Zoological Museum of Moscow University, Russia Preparation techniques and terminology External characters were examined and pencil drawings or photographs were produced using a Wild M10 dissecting binocular microscope with a drawing tube or a digital camera attached to it (courtesy of Muséum d’histoire naturelle, Genève and SEL ARS USDA, c/o USNM). Figures of terminalia (macerated in 10% potassium hydroxide and washed in distilled water) were made from temporary slides in glycerol under a Wild M11-36976 compound binocular microscope, with a camera lucida. Photographs of wings were made from dry slides with detached softened wings. After examination the detached parts were placed in gelatin capsules or plastic tubes with glycerol, closed with a stopper, and pinned beneath the respective specimens. Series of digital images of three-dimensional structures were combined to produce sharper pictures of whole bodies and heads. Pencil drafts were redrawn using Genius NewSketch 1812D drawing pad, and compound computer image processing and drawing techniques in Adobe® Photoshop® 6.0. Morphological terminology was generally adopted from White et al. (1999). Cladistic analysis The program TreeGardener 2.2 (Ramos, 1997) was used to prepare the character matrix containing thirty-two characters and 26 taxa, including 25 species of Myennidini plus the outgroup, which was scored with what we believe to be the plesiomorphic state for each character (or for characters of uncertain polarity, as unknown (?)). The matrix was prepared in HENNIG86 format (Table 1), and was then converted to NEXUS format and edited using NEXUS Data Editor (Page, 2001a). These morphological data were then analyzed using the programs HENNIG86 (Farris, 1988) and PAUP 4.0 beta 4a version for Windows (Swofford, 1998). Further details regarding the options used in the analysis are explained below, in the ‘Discussion of the phylogenetic analysis’. The resulting trees were saved and imported into the program TreeView (Win32) 1.6.5 (Page, 2001b) to produce vector tree plots. This program was also used to save tree files in a format compatible with PAUP. Trees obtained from HENNIG86 were also imported into PAUP 4.0 to trace their character state changes. The first co-author (EPK) is responsible for the results and conclusions of this work in general. The second co-author (VAK) prepared the digital illustrations and provided computer phylogenetic analysis.

1. CLASSIFICATION OF ULIDIIDAE

Family Ulidiidae Ulidini Macquart, 1835: 498. Ulidioidae Agassiz, 1846: 383. Ulididae Bigot, 1852: 488. Ulidiinae Bezzi, 1894: 332. Ulidiidae Sharp, 1899: 504; Hendel, 1936a: 77; 1936b: 1947; Hennig, 1940: 1; 1952: 221; 1973: 54; Brues et al., 1954: 369; Zuska, 1967: 201; Richter, 1970b: 130; Mamaev et al., 1977: 247; Zaitzev, 1982: 422; 1984: 59; Krivosheina and Krivosheina, 1995a : 49, 1995b: 109, 1997a: 460, 1997 d: 1179; Merz, 1996b: 405; 1998: 242; Clements and Merz, 1997: 65; 1998: 55; Clements, 1998: 122; Greve, 1998: 187;

500 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Korneyev, 1999: 12; Barraclough, 2000: 77. For other spellings see synonymy of the tribe Ulidiini below. Ortalidae: Curran, 1934a: 420. Otitidae Aldrich, 1932: 7; Malloch, 1932: 205, 1933: 247, 1940: 205; Curran, 1934b: 271; Hendel, 1936b: 1947; Harriot, 1942a: 23; 1942b: 195; Hennig, 1952: 220; 1958: 578; 1973: 16, 18, 55; Brues et al., 1954: 371; Steyskal, 1961: 401, 403; 1965: 643; 1968: 54.1; 1977: 165; 1980: 575; Richter, 1970a: 123; Mamaev et al., 1977: 247; Soós, 1984: 45; Evenhuis, 1989: 479; Thompson and Pont, 1994: 163; Krivosheina and Kovalev, 1972: 81; Merz, 1996a: 329; Krivosheina and Krivosheina, 1997b: 628, 1997c: 671; Greve, 1998: 185, 187. Pterocallidae: Hendel, 1916: 297; 1936b: 1946; Brues et al., 1954: 370. Diagnosis Diversely colored, usually picture-winged, acalyptrate possessing most characters of the superfamily Tephritoidea: 1) tibiae without dorsal row of setae; 2) surstylus with prensisetae (= thickened setae) mesally; 3) phallus long and coiled; 4) well-developed telescopic ovipositor consisting of oviscape (= fused tergosternum 7), eversible membrane bearing 2 pairs of taenia impregnated by chitin, and aculeus consisting of 4 longitudinal sclerites, paired derivatives of tergite and sternite 8, and apical cap-like cercal unit with sensory setae. The Ulidiidae differ from other families of Tephritoidea by the following combination of characters: fronto-orbital plates not developed and not bearing strong setae (except in Loew and allied genera); presutural supra-alar (except in Dyscrasis Hendel and some species of Otites Latreille) and katepimeral setae lacking (presutural supra-alar present in the ground plan of most other families; katepimeral setae or setulae present in Platystomatidae, Ctenostylidae, Pyrgotidae and Tephritidae); vein Sc entire and slightly bowed, neither bent anteriorly at right angle, nor constricted before apex (bent anteriorly at right angle in Pyrgotidae Toxurini and most Tephritidae, and constricted or broken before apex in Ctenostylidae, most Tephritidae and some

Pyrgotidae); vein R1 usually setulose only on apical portion (except in all Ulidiini and most Lipsanini bare, and certain species of other tribes sometimes either bare or entirely setulose) (bare in most or all Piophilidae, Pallopteridae, Lonchaeidae and Richardiidae, and entirely setulose in Platystomatidae, Ctenostylidae, Pyrgotidae and Tephritidae); cell bcu with triangular posterodistal lobe, except in Zetterstedt and a few Pterocallini (absent in lower Tephritoidea, all Platystomatidae and Ctenostylidae and a few genera of Pyrgotidae and Tephritidae); male tergite 6 lacking (rudimentary in a few Lonchaeidae and Pallopteridae); male sternite 6 bare (setulose in lower Tephritoidea); abdominal spiracles 6 and 7 lacking in males (present in many lower Tephritoidea); phallus, when at rest, stored in pouch on ventral aspect of abdomen; phallus apex not forming separated glans (stored on dorsal side underneath tergite 6 and bearing glans in other higher Tephritoidea); taeniae of eversible membrane bare, bearing no trichoid sensilla (bearing trichoid sensilla along whole length in most Piophilidae, Pallopteridae and Lonchaeidae) and having 2 ducts of spermathecae, one of them apically bifurcated and bearing 2 spermathecae, except Lipsanini, which have 2 spermathecae on 2 ducts (3 spermathecal ducts in ground plan of Pyrgotidae and Tephritidae; 2 of them bifurcated close to vagina). Ground plan set of setae: 2 orbital (anterior seta shorter and usually in front of margin of vertical plate); medial and lateral vertical well-developed; 1 postpronotal; 1 anepisternal (at ventral margin), setulae in dorsal half of anepisternum weak; 1 acrostichal; 2 postsutural dorsocentral setae, anterior aligned with, or slightly posterior to, postsutural supra-alar setae,

501 BIOTAXONOMY OF TEPHRITOIDEA posterior aligned with intra-alar setae, lateral to anterior dorsocentral seta; 1 postsutural supra- alar, 1 intra-alar and 1 postalar; 2 pairs of scutellars; 1 or 2 postphragmal anepisternal and 1 katepisternal. Included subfamilies Ulidiinae and Otitinae.

Subfamily Ulidiinae Macquart, 1835 Ulidini Macquart, 1835: 498. Ulidiinae Bezzi, 1894: 332; Becker, 1905: 105; Hendel, 1909d: 151; 1910b: 1, 1912: 5; Hennig, 1940: 10; 1952: 222; 1958: 578; 1973: 16; 54; Steyskal, 1952: 277; 1961: 403; 1965: 650; 1968: 54.13; 1977: 166; 1980: 575; Brues et al., 1954: 369; Evenhuis, 1989: 480. For other spellings see synonymy of the tribe Ulidiini below. Diagnosis Flies of diverse appearance, variously colored, from yellow to black and from dull gray to shiny green, with vein R1 setulose or bare, and phallus bare (rarely trichose or spinulose); female abdominal sternites 4–6 lacking anterior apodemes. Discussion Hennig (1940, 1958, 1973) and Steyskal (1961, 1965, 1968, 1987) restricted the concept of

Ulidiinae to include only genera with vein R1 and phallus bare. Based on other characters, we found that some genera with phallus and vein R1 setulose are closely related to other genera that have them bare (Kameneva and Korneyev, 1994, Kameneva and Korneyev in Korneyev, 1999, and unpublished data). The setulae on both vein R1 and the phallus are now considered as characters highly subject to homoplasy and correlated. So far, the only hypothesized apomorphy of the subfamily is the absence of the anterior apodemes on the female sternites 4– 6. The subfamily contains the following four tribes, Seiopterini (predominantly Holarctic), Ulidiini (Old World) and Pterocallini and Lipsanini (predominantly Neotropical tribes). Tribe Ulidiini Macquart, 1835 Ulidini Macquart, 1835: 498. Ulidiina Hendel, 1910a: 104. Ulidiini Kameneva and Korneyev, 1994: 68. Other spellings: Ulidiai Desmarest, 1849: 753; Uliditae Desmarest, 1860: 67; Uliditi Lioy, 1864: 1092; Ulidina Loew, 1868a: 283, 1873: 64; Ulidiides Rye, 1873: 395; Ulidi Bigot, 1886: 289; Ulidiae Bigot, 1892: 227. Type genus: Meigen, 1826: 385. Type species: Ulidia erythrophthalma Meigen, 1826 (designated by Hennig, 1940: 13). Diagnosis Black or reddish-yellow flies, sometimes with metallic green or blue sheen (in Fallén and some species of Wiedemann), with moderately or very robust body and predominantly hyaline wings, often with subbasal and apical dark spots. Vein R1 bare, cell r4+5 more or less narrowed towards apex; chaetotaxy unmodified relative to family ground plan (see above). Male terminalia: surstylus usually with several prensisetae, phallus guide (longitudinal depression of hypandrium anterior of basiphallus, between gonites) shallow and unmodified (plesiomorphies); phallus large, thick and mostly bare, with apical area bearing several sclerotised hook-like appendages, similar to glans of Tephritidae (apomorphy). Female terminalia: tergite and sternite 6 strongly narrowed and partially fused forming a narrow ring tightly constricting the base of oviscape (see: Kameneva, 1996) (apomorphy); aculeus unmodified relative to family ground plan, in a few species of Timia Wiedemann with apical

502 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) unit acute and fused to tergite 8; 3 round spermathecae, smooth or covered (in Physiphora) by acutely pointed papillae. Genera included The tribe comprises 3 Old World genera. Physiphora is predominantly African, with a few endemic species in the Oriental Region and Solomon Islands, and some synanthropic species introduced into the New World and Australasia, whereas Ulidia Meigen and Timia are most abundant in all semiarid and arid areas of the Palaearctic Region. Discussion The monophyly of the tribe is well supported by the shape of the phallus tip (see Hennig, 1940: Textfig. 2–3) in combination with a bare or short spinulose distiphallus, and modified female tergite and sternite 6. Certain problems relevant to nomenclature of the tribe still need to be resolved: 1. The current concept of the type genus of the tribe, which was broadly accepted in common use during the 20th century, is based upon an erroneous Hendel’s (1910a) statement that Ulidia Meigen, 1826 was established as a separate genus rather than a replacement name for Chrysomyza Fallén, 1817 (= Physiphora Fallén, 1817). Coquillett (1911) was the first to note that: “Hendel is in error in stating that Ulidia is not a synonym of Physiphora. When treating of this genus, Meigen expressly states that Fallén named it Chrysomyza (a change of name for Physiphora under the mistaken impression that it was preoccupied) and that he (Fallén) knew only one species. But as this name was not appropriate for the two new species that Meigen added, he tells us that he changed the name to Ulidia (Syst. Beschr. vol. V, p. 386). Thus Meigen was not justified in changing the name previously given to this genus by Fallén, and it follows that Ulidia must be a synonym of Physiphora”. This important comment was ignored, and the subsequent use of the name Ulidia sensu Hendel was maintained and came into common use. Hennig (1940: 45) designated Ulidia erythrophthalma Meigen, 1826 as the type species of Ulidia Meigen, 1826, and this designation was accepted as valid (e.g., Zaitzev, 1984). The name Ulidia, prevalent during the 1800s and 1900s, was sensu Hendel and Hennig. Sabrosky (1999) noted in his discussion of the family-group name Ulidiidae: “I agree with Coquillett (1910: 618), who stated that Ulidia was a change of name for Chrysomyza Fallén, 1817 “on the ground that the latter name is not applicable to all the species” (Meigen: “auf die andern nicht passt”). As a change of name, the type would be Chrysomyza demandata (Fabricius), automatically [demandata was included in Ulidia Meigen]. If it were a new proposal for part of the species, it would require its own type species, the view taken by Hennig (1940). If the name Ulidia had fallen as a synonym of Physiphora, q.v. (synonym Chrysomyza, q.v.), the species of Ulidia would have been left without a generic name, there being no synonyms. Recognition of Ulidia would in my opinion require action by the Commission.” Following Sabrosky’s opinion, we maintain Hennig’s designation of Ulidia erythrophthalma Meigen, 1826. According to Articles 65.2 and 70.2 (ICZN, 1999), this case should be referred to the Commission for a ruling, as the universality of use of the name Ulidia is threatened. 2. Nomenclatural stability is threatened also by the fact that Timia Wiedemann, 1824 and Ulidia Meigen, 1826 are two closely-related genera, and if the forthcoming revision of these genera (Kameneva, in prep.) shows that there is no way to recognize them as separate genera, then Ulidia, the family-name bearer, could become a junior subjective synonym. However, neither such a nomenclatural action, nor the possible synonymy with Physiphora

503 BIOTAXONOMY OF TEPHRITOIDEA discussed above, would affect the validity of the family group names Ulidiini, Ulidiinae and Ulidiidae (ICZN, 1999: Art. 40.1). Tribe Seiopterini Kameneva and Korneyev, 1994 Myodinae Robineau-Desvoidy, 1826: 17 (unavailable; prior to available generic name; in a suppressed work). Myodinae Robineau-Desvoidy, 1830: 704; corrected spelling: Myodininae (invalid name based on a junior synonym and never widely accepted); other spellings: Myiodinae Agassiz, 1846: 242. Type genus: Myodina Robineau-Desvoidy, 1830: 727. Type species: Musca urticae Fabricius, 1794 (by monotypy; type species misidentified, teste Macquart, 1835: 437) = Musca vibrans Linnaeus, 1758 (now in Seioptera). Ortalides Fallén, 1810: 15. Ortalidae Swainson, 1840: 378. Ortalididae Harris, 1841: 416. Ortalidiformes Frey, 1921: 216, 217. Other spellings: Ortalideae Macquart, 1835: 429; Ortalida Heer, 1849: 252; Ortalidina Rondani, 1856: 24, 108; Ortaliditae Desmarest, 1860: 45; Ortaliti Lioy, 1864: 1017; Ortalidae Loew, 1866: 238; Orthalidinae Anonymous, 1876: 180; Orthalidae Osten Sacken, 1881a, 1881b: xcix; Ortalidi Bigot, 1886: 289; Orthalididae Beschovski, 1972: 10. Type genus: Ortalis Fallén, 1810: 17 (preoccupied: Merrem, 1786 in Aves) [stem Ortalid-]. Type species: Musca vibrans Linnaeus, 1758 (designated by Westwood, 1840b: 149) (now in Seioptera). Valid name: Seioptera Kirby, 1817. Junior synonym: Ortaliscus Ghesquière, 1947: 691. Ortaliscidae Ghesquière, 1947: 691 (invalid name based on a junior synonym and never widely accepted). Type genus: Ortaliscus Ghesquière, 1947: 691 (new name for Ortalis Fallén). Type species: Musca vibrans Linnaeus, 1758 (automatic) (now in Seioptera). Valid name: Seioptera Kirby, 1817. Seiopteridae Sabrosky, 1946: 170 [unavailable; suggested in discussion of a family-name problem, but not adopted]. Seiopterini Kameneva and Korneyev, 1994: 68, 69. Type genus: Seioptera Kirby, 1817: 305. Type species: Musca vibrans Linnaeus, 1758 (by monotypy, cited as “Tephritis vibrans, Latr.”). Preoccupied senior synonym: Ortalis Fallén, 1810. Junior synonym: Myodina Robineau-Desvoidy, 1830. Diagnosis Shiny black flies with 2 katepisternal setae, 4 spherical spermathecae, and anepisternal setae lacking (synapomorphies of the tribe). Vein R1 entirely setulose (Pseudoseioptera Stackelberg and Homalocephala angustata Wahlberg) or bare, phallus either bare or short setulose. Genera included The tribe is comprised of 3 holarctic genera. Homalocephala Zetterstedt, 1839 and Pseudoseioptera Stackelberg, 1955 are both associated with pine and mixed forests; their larvae live under the bark of fallen trees (Krivosheina and Kovalev, 1973; Krivosheina and Krivosheina, 1995 b). The third genus, Seioptera Kirby, 1817, includes a synanthropic species (Linnaeus) whose larvae live in decomposing plant matter and dung. Discussion The monophyly and concept of this tribe were delimited by Kameneva and Korneyev (1994), and later Kameneva and Korneyev in Korneyev (1999) showed that it belongs to the Ulidiinae because of the absence of the sternal apodemes in females.

504 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Tribe Pterocallini Loew, 1868 Pterocallina Loew, 1868b: 1, 8; 1873: 61; Becker, 1905: 104. Pterocallinae Snow, 1896: 115, 117; Hendel, 1909a: 6, 1909b: 1; Aldrich, 1932: 7; Aczél, 1951: 397; Hennig, 1952: 221, 1958: 578; 1973: 55. Pterocallidae Hendel, 1916: 297, 1936b: 1946; Brues et al., 1954: 370. Pterocallini Kameneva and Korneyev, 1994: 68. — Perocall.[-inae] Hendel, 1914b: 172 [error]. Type genus: Pterocalla Rondani, 1848: 83. Type species: Dictya ocellata Fabricius, 1805 (by monotypy). Diagnosis

Dull brown or gray flies. Vein R1 setulose or bare. Most members sexually dimorphic: in males, pterostigma usually strongly enlarged (as long as, or even longer than costal cell) and vein R2+3 usually sinuous (if sometimes straight, then vein R4+5 setulose) (apomorphies). Male terminalia: the part of the ejaculatory apodeme adjoining the sperm sac elongate, more than twice as long as wide (synapomorphy with Lipsanini?); phallus bare or short sparsely setulose. Female terminalia: 3 spherical spermathecae, with smooth surface. Genera included The tribe contains 23 neotropical genera: Aciuroides Hendel, 1914b, Bothrometopa Hendel, 1909a, Chondrometopum Hendel, 1909a, Coscinum Hendel, 1909a, Cymatozus Enderlein, 1912b, Cyrtomostoma Hendel, 1909a, Dasymetopa Loew, 1868 (= Euxestina Curran, 1934a), Elapata Hendel, 1909a, Goniaeola Hendel, 1909a, Idanophana Hering, 1938, Megalaemyia Hendel, 1909a, Micropterocerus Hendel, 1914, Neomyennis Hendel, 1914c, Ophthalmoptera Hendel, 1909a, Paragorgopis Giglio-Tos, 1893, Wiedemann, 1830, Pterocalla Rondani, 1848 (including Pseudopterocalla Hendel, 1909a), Pterocerina Hendel, 1909a, Rhyparella Hendel, 1909a, Sympaectria Hendel, 1909a, Terpnomyennis Kameneva, 2004, Terpnomyia Hendel, 1909a, Tetrapleura Schiner, 1868 and Xanthacrona Wulp, 1899. The genus Schnusimyia Hendel, 1914a should be transferred to Richardiidae based upon the presence of well-developed spiracles 6 and 7 and setulose abdominal sternite 6 in the male (Norrbom, unpublished data; Korneyev, unpublished data). Neomyennis and Xanthacrona, often associated with the Myennis group of Otitinae (Steyskal, 1982, 1987) do not have the unpaired lobe between surstyli (“metaphallic plate”) that characterizes the latter group, but absence of female sternal apodemes, possession of 3 round spermathecae and male terminalia similar to those in Pterocalla clearly puts them here, in Ulidiinae, not Otitinae. Discussion Monophyly of the tribe is supported by the extremely elongate part of the ejaculatory apodeme adjoining the sperm sac; this part usually exceeds the length of the fan-like part, and by the sinuous vein R2+3, a character that may be highly subject to homoplasy. Male genital characters appear rather uniform in each group of genera, similar to Pterocalla, Dasymetopa and Pterocerina, but they still need to be thoroughly examined in most genera of Pterocallini in order to clarify the limits of the genera and the relationships among them. Tribe Lipsanini Enderlein, 1938 Lipsaninae Enderlein, 1938: 676; Evenhuis, 1989: 481; Sabrosky, 1999: 179. Lipsanini: Kameneva and Korneyev, 1994: 68. Type genus: Lipsana Enderlein, 1938: 676. Type species: L. insulaepaschalis Enderlein, 1938 (orig. des.). Euxestinae Hennig, 1940: 3, 4; 1952: 222; 1958: 578; 1973: 16, 54; Zaitzev, 1984: 64;

505 BIOTAXONOMY OF TEPHRITOIDEA

Kameneva, 1992: 24. Euxestidae Frey, 1964: 3; Sabrosky, 1999: 137 (preoccupied by Euxestinae Grouvelle, 1908 in Coleoptera). Type genus: Euxesta Loew, 1868a: 297. Type species: Ortalis notata Wiedemann, 1830 (designated by Coquillett, 1910: 543). Chaetopsidae Crampton, 1944: 153. Type genus: Chaetopsis Loew, 1868a: 315. Type species: Ortalis aenea Wiedemann, 1830 (designated by Coquillett, 1910: 521). Diagnosis Usually metallic green or bluish, often sparsely microtrichose, occasionally shiny or dull black, or completely gray to brown microtrichose flies. Vein R1 bare or, in a few species of Pareuxesta Coquillett and Chaetopsis Loew, setulose along extreme apex. Wing venation not sexually dimorphic; pterostigma not enlarged (shorter than costal cell) and vein R2+3 usually straight. Male terminalia different in two main groups. In (the type species), margins of phallic guide (fovea of hypandrium anterior to basiphallus) not strongly produced; surstylus bearing 2-4 prensisetae. In most other examined genera, including some species currently placed in Euxesta, phallic guide deep, with highly projected lateral lobes; epandrium very short, without trace of medial surstylus and prensisetae. Part of ejaculatory apodeme adjoining sperm sac usually elongate, about twice as long as wide (synapomorphy with Pterocallini?). Phallus bare or, rarely, setulose (in Loew, 1868a) (see Steyskal, 1963a) or even spinulose (in Euacaina Steyskal, 1963). Female terminalia: 2 spherical spermathecae (synapomorphy), usually with smooth surface or, in some genera, covered by acute papillae. Female tergite 6 well developed (plesiomorphy) at least in Eumetopiella Hendel, and a few allied genera, shortened and narrowed in the majority of genera. Genera included Loew, 1868; Acrostictomyia Blanchard, 1938; Aspistomella Hendel, 1909; Axiologina Hendel, 1909; Cenchrometopa Hendel, 1909; Chaetopsis Loew, 1868; Euacaina Steyskal, 1963; Eumecosomyia Hendel, 1909; Eumetopiella Hendel, 1907; Euphara Loew, 1868; Euxesta Loew, 1868a (= Amethysa Macquart, 1835: see discussion below; Aloceuxesta Hendel, 1936; Euxestina Enderlein, 1937, nec Curran, 1934a); Heterodoxa Malloch, 1932; Hypoecta Loew, 1868a; Lipsana Enderlein, 1938; Notogramma Loew, 1868; Paraphyola Hendel, 1909; Pareuxesta Coquillett, 1901; Malloch, 1932; Polyteloptera Hendel, 1909; Pseudeuxesta Hendel, 1910; Siopa Hendel, 1909; Stenomyia Loew, 1868; Ulivellia Speiser, 1929; Vladolinia Kameneva, 2005; Zacompsia Coquillett, 1901 (= Metopocampta Enderlein, 1927). Discussion The synapomorphy of the genera examined (Hypoecta, Polyteloptera, Lipsana and Acrostictomyia species were not dissected), which substantiates the monophyly of the tribe, is apparently the presence of two spermathecae. The relationships among the genera are not well understood. The largest genus of the tribe, Euxesta, which was defined mainly by the absence of characters of the other genera, is heterogeneous and probably not monophyletic. A revision of that genus, including phylogenetic analysis, may clarify its limits. Barraclough (1999) recently redescribed the type species of Amethysa Macquart — A. fasciata Macquart, 1835 from Cuba — which seems to be very close to, or identical with, E. annonae (Fabricius). To prevent nomenclatural instability, it is strongly recommended not to use the name Amethysa rather than the widely-used name Euxesta, at least until a revision of the latter has been carried out. The tribal name Lipsanini has priority over both Euxestini Hennig, 1940 and Chaetopsini

506 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Crampton, 1944. However Lipsana Enderlein, 1938 is close to, and may eventually become a junior synonym of either Euxesta Loew, Neoeuxesta Malloch or Heterodoxa Malloch, although the type specimens of Lipsana insulaepaschalis Enderlein, 1938 have not been located, and this synonymy is still uncertain. According to Sabrosky (1999), the tribal name Euxestini is unavailable. The family-group name based on Euxesta is preoccupied by Euxestinae Grouvelle, 1908, based on Euxestus Wollaston, 1848 (Coleoptera: Cerylonidae). The name Euxestidae was cited in the synonymy of Cerylonidae (Lawrence, 1982; as cited by Sabrosky, 1999). The third name, Chaetopsini, is based on a valid genus name and is available. It might be validated by the suppression of Lipsanini.

Subfamily Otitinae Aldrich, 1932 Aldrich, 1932: 7; Hennig, 1952: 222; 1958: 578; Steyskal, 1961: 401, 403; 1965: 643; 1968: 54.1; 1977: 165; 1980: 575; Evenhuis, 1989: 479. Diagnosis

Mostly dull gray to shiny brown or black flies with vein R1 setulose or, rarely, bare, and a spinulose (rarely bare) phallus, and with the female abdominal sternites 4–6 with anterior apodemes. Discussion The concept of the Otitinae adopted by Hennig (1939, 1958; as Otitidae) and Steyskal (1961,

1965, 1968, 1987) was much broader, including most Ulidiidae genera with setulose vein R1 and phallus. After reassigning the holarctic Seiopterini and neotropical Pterocallini (both including genera with bare and setulose R1) to the Ulidiinae, the subfamily is now restricted to genera with a spinulose phallus (synapomorphy?) and the female sternal apodemes present (symplesiomorphy?). It can be subdivided into the following three tribes: Cephaliini, Otitini and Myennidini, new tribus. Tribe Cephaliini Schiner, 1864 Cephalinae Schiner, 1864: 63. Cephaliinae Snow, 1896: 115, 116; Sabrosky, 1999: 77. Cephaliidae Sturtevant, 1926: 213. Other spellings: Cephalina Loew, 1868b: 1, 7; Cephalini Kameneva and Korneyev, 1994: 65, 68. Type genus: Meigen, 1826: 293. Type species: C. nigripes Meigen, 1826 (by subsequent designation of E. Blanchard, 1846: pl. 181) = C. rufipes Meigen, 1826. Junior synonym: Myrmecomya Robineau-Desvoidy, 1830. Myrmecomyiinae Hendel, 1910c: 310; 1914: 158; 1936b: 1945; Sabrosky, 1999: 208. Other spellings: Myrmecomyinae Hendel, 1912: 2. Type genus: Myrmecomya Robineau- Desvoidy, 1830: 721. Type species: Myrmecomya formicaria Robineau-Desvoidy, 1830 (designated by Coquillett, 1910: 573) = Cephalia rufipes Meigen, 1826. Diagnosis Palpus large, broad, subtriangular; clypeus often large, straight in profile; ventral facial margin often strongly produced; arista short pubescent or bare. Proepisternal seta often minute or lacking; scutal setulae often divided by 2–5 bare, finely microtrichose stripes (synapomorphies?).

507 BIOTAXONOMY OF TEPHRITOIDEA

Wing. Vein R1 setulose (completely or on apical portion only); vein R4+5 bare, except setulose dorsally in Acrostictella Hendel. Male terminalia. 2-12 prensisetae well developed; phallus spinulose. Female terminalia. cercal unit unmodified; 3 bare or sparsely papillose, spherical or subglobose spermathecae (plesiomorphies?). Flies of several genera of this tribe have a very characteristic ant-like appearance. Discussion The tribe comprises mostly American genera, with a few representatives in the Palaearctic Region. Genera included Acrostictella Hendel, 1914; Cephalia Meigen, 1826; Delphinia Robineau-Desvoidy, 1830; Myiomyrmica Steyskal, 1961; Myrmecothea Hendel, 1910; Proterpnomyia Blanchard, 1967; Proteseia Korneyev and Hernández, 1999; Pterotaenia Rondani, 1868, genus resurrected from synonymy; Loew, 1873. The species of the genus Pterotaenia were reexamined and found to be members of the Cephaliini rather than Ceroxys Macquart (Otitini). These two genera have an acute first flagellomere and a banded wing pattern, but are otherwise different. Further, most of the species assigned to the genus Terpnomyia Hendel, 1909, except its type species (which belongs in Ulidiinae Pterocallini), should actually be transferred to the Proterpnomyia. The monophyly of the tribe is supported by a single synapomorphy (palpus extremely broad), plus possibly by the presence of bare mesonotal stripes (synapomorphy? shared with the following group) in combination with some plesiomorphic characters (apodemes of abdominal sternites 4-6 well developed; spherical spermathecae). However, a more detailed study of the genera discussed above is beyond the scope of this paper. Genera incertae sedis, possibly related to Cephaliini A group of genera, which have microtrichose non-setulose vittae on the mesonotum but do not possess broad subtriangular palpus, and are possibly related to the Cephaliini rather than the Otitini, includes the North American Diacrita Gerstäcker, 1860, Loew, 1868, and the Andean Pseudomelieria Brèthes, 1921. The nearctic genus Psaeropterella Hendel, 1914 and its relatives, Curranops Harriot, 1942; Haigia Steyskal, 1961 and Steyskal, 1961, may also be related to the Cephaliini, sharing similar epandrial characters with Cephalia, Diacrita and Idana. Tribe Otitini Aldrich, 1932 Otitides Westwood, 1840: 146 (unavailable, not adopted). Tetanopina Loew, 1862: 40. Type genus: Tetanops Fallén, 1820. Type species: T. myopina Fallén, 1820 (by monotypy) = T. myopinus Fallén. Otitidae Aldrich, 1932: 7. Otitiformes and Otitoidea Crampton, 1944: 153. Otidimorpha Brues et al., 1954: 22. Other spellings: Otididae Hardy, 1959: 229; Otitiidae Rohdendorf, 1964: 17. Type genus. Otites Latreille, 1804. Type species: O. porca Latreille, 1804 (by monotypy; as “Musca porcus Bosc,” a manuscript name validated there by Latreille).

508 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Ceroxydidae Steyskal, 1946: 168 (nomen nudum; not diagnosed and not adopted). Type genus. Ceroxys Macquart, 1835. Type species: Musca urticae Linnaeus, 1758 (designated by Westwood, 1840: 149). Diagnosis Palpus usually moderately broad or linear, but not extremely broad; nor subtriangular (except in oscillans Meigen); clypeus low, slightly convex in profile; face usually with carina and antennal grooves; ventral facial margin not strongly produced; arista short or long pubescent; proepisternal seta well developed. Scutal setulae irregular or arranged in rows, not separated by microtrichose or bare vittae. Male terminalia: prensisetae well developed, mediodorsal (paracercal) prensiseta either present or absent; paired sclerites of epiphallus usually developed posterior of basiphallus (except in Herina); unpaired metaphallic lobe not developed between bases of surstyli (see description of this structure under Myennidini); phallus spinulose. Female terminalia: cercal unit usually unmodified; 3 elongate, wrinkled or papillose spermathecae, very rarely smooth and spherical or subglobose. Genera included The tribe is distributed in the Old World and North America, and is absent in South America and the temperate part of the Australasian Region. It includes the widespread genera Herina Robineau-Desvoidy (predominantly palaearctic, with some nearctic and caribbean, oriental and australasian species assigned to it) and Robineau-Desvoidy (predominantly holarctic, with at least one species in Africa), and the holarctic genera Otites Latreille, Tetanops Fallén, and Ceroxys Macquart. The tribe also includes the nearctic genus Hiatus Cresson and the strictly palaearctic genera Meigen and Ulidiopsis Hennig. Discussion Recently, all species assigned to the palaearctic genera Loew and Becker were placed in Melieria (Kameneva, 1997a), and the Palaearctic species of Systata Loew were transferred to Otites (Kameneva, 1997b). Dorycera Meigen and Ulidiopsis Hennig include species that are very closely related to the genera Otites and Herina Robineau- Desvoidy, respectively, and could be considered aberrant members of them. However, the concepts of the two latter genera also need further revision (Kameneva, pers. observation; Merz, pers. comm.). Some American genera previously assigned to the Otitinae (Steyskal, 1961, 1965, 1968) are tentatively associated here with the Cephaliini (see above). However, the characters that differentiate Otitini from Cephaliini are not sufficiently reliable. The concept of the tribe Otitini and its composition therefore need special reconsideration. Tribe Myennidini Kameneva and Korneyev, new tribe Type genus: Myennis Robineau-Desvoidy, 1830: 717. Type species: Scatophaga fasciata Fabricius, 1805 (by monotypy) = Musca octopunctata Coquebert, 1798. Diagnosis Face flattened, without or with low (in Pseudodyscrasis and Namibotites) medial carina; antennal groove indistinct, except in Namibotites; clypeus moderately developed, convex in profile (in Pseudotephritis, Callopistromyia, Pseudotephritina and Myennis) or small, but not high and straight in profile; ventral facial margin not strongly produced. Arista bare. Palpus not

509 BIOTAXONOMY OF TEPHRITOIDEA subtriangular, usually moderately large or, in Dyscrasis and Pseudodyscrasis, very large.

Prosternum non-setulose and microtrichose, in one species setulose. Vein R1 setulose, except in Oedopa, Paroedopa, Stictomyia and Ulidiotites. Proepisternal seta moderately developed or minute; scutal setulae usually irregular, except well-expressed dorsocentral row. Male terminalia. paired sclerites of epiphallus posterior to basiphallus absent; unpaired metaphallic lobe developed between bases of surstyli (apomorphy); phallus spinulose, setulose or bare. Female terminalia. tergosternum 8 (aculeus basal portion) well-sclerotized, long and narrow (apomorphy?), cercal unit unmodified; 3 smooth, spherical or subglobose spermathecae (plesiomorphies?), except in Namibotites. Genera included This tribe is predominantly holarctic (Appendix 3). It comprises the palaearctic genus Myennis Robineau-Desvoidy, 1830, the nearctic, or predominantly nearctic genera Oedopa Loew, 1868, Stictomyia Bigot, 1885, Paroedopa Coquillett, 1900, Callopistromyia Hendel, 1907 (= Callopistria Loew, 1873 (preoccupied by Callopistria Hübner, 1816)), Pseudotephritina Malloch, 1931, Dyscrasis Aldrich, 1932, Ulidiotites Steyskal, 1961, the holarctic Pseudotephritis Johnson, 1902 (= Stictocephala Loew, 1873 (preoccupied by Stictocephala Stål, 1869)), the mainly Mesoamerican Pseudodyscrasis Hernández, 1988, Neodyscrasis Kameneva and Korneyev, n. gen., South American Stictoedopa Brèthes, 1926, and the South African Arborotites Barraclough, 2000 and Namibotites Barraclough, 2000. Discussion The genera placed here were previously assigned to the “Pterocallidae” (Hendel, 1909b; Hennig, 1939) because of their dull grayish, often brown spotted, appearance and lack of a facial carina and antennal grooves. Callopistromyia strigula (Loew, 1873) was classified in the genus Pterocalla Rondani (Pterocallini) because of its elongate wing shape and the spotted pattern characteristic of the latter genus. The terminalia characters in both tribes, however, are very distinctive (Steyskal, 1979), and they belong to different subfamilies. The monophyly of the new tribe is supported by at least one unique synapomorphy not known elsewhere in the Tephritoidea: an area posterior to the basiphallus forms a flattened two- walled metaphallic lobe often bearing a few sparse and small setulae (e.g., Figs. 12, 16, 26, 27, 36). From its position, this structure is probably associated with the proctiger, rather than with the basiphallus; in Ulidiotites it appears to be an anterior flap of the proctiger. Various genera, such as Myennis, Dyscrasis and Oedopa, share this character in combination with the presence of anterior apodemes of female sternites (3-)4-6. In contrast to other Myennidini, Oedopa,

Paroedopa, Stictomyia, Ulidiotites and Namibotites have vein R1 bare and the phallus non- spinulose, both characters previously believed to be characteristic of the Ulidiinae (Hennig, 1958, 1973; Steyskal, 1961, 1987). The current study shows that both these characters are inconsistent and are subject to homoplastic changes in three different lineages of Ulidiinae (see above), as well as in this tribe (see the phylogenetic analysis below).

510 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

2. REVIEW OF MYENNIDINI GENERA

Key to genera of Myennidini

1. Vein R1 entirely bare ...... 2

- Vein R1 dorsally setulose at least apically ...... 8 2. First flagellomere more than 3 times as long as wide. Wings dark brown to gray, with reticulate pattern of hyaline spots (Figs. 50, 56). North America ...... Stictomyia Bigot - First flagellomere short oval, at most twice as long as wide. Wings predominantly hyaline, sometimes with few brown spots on apical half ...... 3 3. Face between antennae and frons with dark brown or black spots or transverse bar (Figs. 32, 40, 45). Wing always with pattern of dark spots anteroapically; apical section of vein M almost straight, ending at wing apex (Figs. 35, 46)...... 4 - Face and frons uniformly yellowish gray without dark spots (Figs. 62, 68). Wing hyaline with darkened veins (Fig. 60), or if with dark spots at wing apex and on crossveins, then apical section of vein M strongly bent anteriorly, ending before wing apex (Fig. 49) ...... 5 4. Both face between antennae, and frons with dark brown or black transverse bars (Figs. 32, 40). Crossvein DM-Cu sinuous (Figs. 35, 42). North America ...... Oedopa Loew - Face between antennae with single round spot. Frons with two black spots laterally (Figs. 44, 45). Crossvein DM-Cu straight (if DM-Cu sinuous, see “Oedopa” elegans, couplet 6). North America ...... Paroedopa Coquillett 5. Wing hyaline. Vertex forming ridge, which is almost straight or slightly convex ...... 7 - Wing with dark spots at least at wing apex and on crossveins. Vertical ridge often concave. Central or South America ...... 6 6. Wing with numerous dark spots, more intensive towards base. Crossvein DM-Cu sinuous. Frons with 2 black spots. Vertex shape not examined. Central America ...... “Oedopa” elegans Giglio-Tos - Wing with 5–6 dark spots at wing apex and on crossveins. Crossvein DM-Cu straight. Vertical ridge conspicuously concave between eyes (as in species). South America ...... Stictoedopa Brèthes 7. Extension of cell bcu short, not exceeding level of crossvein BM-Cu (Fig. 60). Antennal groove inconspicuous. Ocellar seta developed. Phallus trichose, coiled, 3-4 times as long as epandrium (Fig. 63). Three round, smooth spermathecae. North America ...... Ulidiotites Steyskal - Extension of cell bcu far exceeding level of crossvein BM-Cu (Fig. 69). Antennal groove deep and short. Ocellar seta completely reduced. Phallus bare, not coiled, 1.5–2 times as long as epandrium (Fig. 70). Two oval, papillose spermathecae (Fig. 73). Southern Africa ...... Namibotites Barraclough 8. Scutellum shiny brown or black, at least with two shiny spots (Figs. 99, 102, 108, 119) .. 9 - Scutellum gray and black microtrichose, without shiny areas (Figs. 23, 76) ...... 13 9. Five dorsocentral, five acrostichal and three supra-alar setae, one of each presutural (Fig.

119). Extension of cell bcu conspicuously longer than vein A1+Cu2. North America ...... Dyscrasis Aldrich

511 BIOTAXONOMY OF TEPHRITOIDEA

- No presutural dorsocentral, acrostichal or supra-alar setae (Figs. 108, 113). Extension of

cell bcu shorter or, at most, as long as vein A1+Cu2 ...... 10 10. Three or four dorsocentral setae; intra-alar seta strong, in both sexes without cluster of fine long setulae around it (Fig. 108, 113). Two equal orbital setae (Fig. 111) ...... 11 - Two or one dorsocentral setae; in females intra-alar seta weak with cluster of fine long setulae around it (Fig. 102). Anterior orbital seta shorter than posterior, or absent ...... 12 11. Scutum without large shiny areas; 1 supra-alar seta. Wing with uniformly darkened basal cells (Fig. 107). North America (Mexico) ...... Pseudodyscrasis Hernández - Scutum with large shiny supra-alar areas; 2 supra-alar setae. Wing basal cells with isolated black dots and streaks and hyaline borders (Fig. 112) ...... Neodyscrasis Kameneva and Korneyev, n. gen. 12. Supra-alar area and lateral portion of tergite 4 shiny black to brown. Middle of wing with 2 oblique crossbands: first crossing from base of costal cell through radial fork to middle of discal cell; second from pterostigma through approximated crossveins R-M and DM-Cu (Fig. 87, 90). Silvery microtrichose areas without round spots at bases of setulae. Palaearctic...... Myennis Robineau-Desvoidy - Supra-alar area and lateral portion of tergite 4 without shiny black spots. Middle of wing with 2 straight crossbands: first crossing from apex of costal cell distad of radial fork to middle of discal cell; second from apex of pterostigma to crossveins R-M and DM-Cu (Fig. 101), often broken between crossveins (Fig. 97). Silvery microtrichose areas broken by numerous round bare spots at bases of setulae. Nearctic species ...... Pseudotephritina Malloch 13. Discal cell with numerous small dark and hyaline spots; its anteroapical angle acute. Wing pattern reticulate or spotted, without crossbands (Figs. 24, 25). Abdominal pleura setulose. Nearctic species ...... Callopistromyia Hendel - Discal cell with extensive areas of solid color, without small spots. Wing pattern consisting of broad, partially broken or confluent crossbands (Figs. 6-9, 77), or if reticulate, then anteroapical angle of discal cell about 90º (Figs. 10-11). Abdominal pleura bare...... 14

14. Extension of cell bcu shorter than or, at most, as long as vein A1+Cu2 (Fig. 6–11). Intra-alar seta strong, in both sexes without cluster of fine long setulae around it; 2 dorsocentral setae (Fig. 3). Nearctic or Palaearctic species ...... Pseudotephritis Johnson

- Extension of cell bcu longer than vein A1+Cu2 (Fig. 77). Intra-alar seta weak, in female almost indistinguishable from cluster of fine setulae around it; 1 dorsocentral seta (Fig. 76). South African species ...... Arborotites Barraclough

Pseudotephritis Johnson, 1902 (Figs. 1-21, 125-127) Stictocephala Loew, 1873: 61, 134 (preoccupied by Stictocephala Stål, 1816, Homoptera). Type species: Ortalis vau Say, 1830 (by original designation of Loew, 1873: 61). Pseudotephritis Johnson, 1902: 144 (new name for Stictocephala Loew). Type species: Ortalis vau Say, 1830 (automatic); Aldrich, 1905: 594; Cresson, 1906: 284; Williston, 1908: 272, 277; Hendel, 1909a: 40; 1909b: 22; Banks, 1914: 138; Malloch, 1931: 1; Curran, 1934b: 279; Aczél, 1951: 411; Steyskal, 1962: 254; 1965: 647; 1975: 148; 1987: 807; Soós, 1984: 58; Krivosheina and Krivosheina, 1997c: 671; Kameneva, 2001: 160.

512 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

No detailed descriptions of this genus in its strict sense have appeared since it was briefly diagnosed by Loew (1873, as Stictocephala), subdivided into two subgenera (Malloch, 1931), partially revised (Steyskal, 1962), and the subgenus Pseudotephritina Malloch removed and elevated to full generic status (Steyskal, 1975, 1987). Hendel (1909b) gave the most detailed description of this genus, but including both Pseudotephritis and Pseudotephritina. Consequently, this genus is redescribed below. Most head measurements given in the redescription for P. vau have same or similar condition in P. approximata and P. millepunctata Hennig. Redescription Medium-sized (3.5–5.5 mm) flies with gray microtrichose body having speckled pattern of brown tomentum without shiny areas or bare dots at bases of setae, and non-setulose abdominal pleura. Head. As wide as thorax, slightly higher than long (1.1 : 1) or as high as long (in P. corticalis Loew), and 1.3 times as wide as high (1.6 in P. corticalis). Eyes very slightly higher than long (1.1 : 1 in P. vau) or longer than high (1.1 : 1 in P. corticalis). Frons more or less protruding anteriorly, in profile forming right or slightly acute angle with dorsal portion of face. Lateral margins almost parallel or slightly convergent anteriorly. Frons white microtrichose, with brown spots and small bare dots at bases of setulae; densely microtrichose median vitta distinct, narrowed anteriorly but not broken by darker spots; posterior third rather uniformly microtrichose, usually without brown spots. One moderately developed lateroclinate and slightly proclinate ocellar seta. Two well developed posterolaterally directed orbital setae; posterior slightly longer than anterior. Frontal setulae black, moderately stout, about 0.25 times as long as orbital setae. Face uniformly white microtrichose, conspicuously concave in profile, with transverse furrow in ventral third and without facial carina or distinct antennal grooves. Clypeus convex, moderately low (in profile about 0.5 as high as first flagellomere width), horse-shoe shaped, in profile as long as palpus and 4-5 times as long as high. Parafacial bare on dorsal 0.16, otherwise microtrichose, broadened on ventral half. Facial ridge bare in P. vau, and with 4–5 setulae in P. corticalis. Anterior part of gena with row of 6–8 upwardly curved setulae not exceeding ventral end of ptilinal suture in P. vau, or 8-10 setae reaching vibrissal edge and facial ridge in P. corticalis. Gena 0.25 (in P. vau) to 0.45 times as high as eye (in P. corticalis). Genal seta well developed. Eye as long as high (in P. vau) or longer than high (in P. corticalis), in live specimens horizontally banded. Vertex rounded, without ridge; postocellar seta divergent, as long as ocellars; medial vertical seta almost twice as long as postocellar; lateral vertical seta 0.7–0.8 times as long as medial seta. Occiput on dorsal portion (= median occipital sclerite, cerebrale) not concave, and slightly swollen on ventral portion (= postgena), with transverse dark mark from dorsal margin of occipital foramen to posterior margin of eye, and more or less distinct brown dots at bases of postocular and occipital setulae. One paravertical setula on each side of median occipital sclerite; 4-7 setulae in single postocular row; occipital setulae forming one oblique row of 3–4 setulae laterally of suture and 4–9 setulae in dorsoventral row along occipital foramen. Anterodorsally curved supracervical setulae in two groups of 5-8 on each side, separated by bare area. Postgena moderately setulose. Hypostomal bridge short and bare. Antennae well separated, distance between their bases 1.7–2.0 times as wide as antennal socket. Scape short. Pedicel wider than long, with blunt triangular medial lobe and shallow dorsal incision; lateral surface with one row of marginal setulae, with 2 or 3 longest

513 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 1–5. Pseudotephritis. 1–2. P. vau. 1. Habitus (from Hendel, 1909b). 2. Occiput. 2a. Palpus, lateral view. 3–4. P. metzi, holotype. 3. Head and mesonotum, dorsal view. 4. Habitus, lateral view. 5. P. corticalis, syntype, head, lateral view (first flagellomere lost). at ventral margin; 12–16 marginal setulae on medial surface forming 1–2 irregular rows; dorsal pedicellar seta almost as long as pedicel. First flagellomere 1.1–1.2 as long as wide, short microtrichose. Two visible basal aristomeres. Terminal aristomere short pubescent. Proboscis stout, with large, rounded prementum that bears 10–12 setae on each side (Fig. 2). Labellum fleshy, large. Palpus (Fig. 2a) rather wide, broadened towards apex, with 12–15 setulae at ventral margin and 5–8 shorter setulae on lateral and medial surfaces. Thorax. Longer than wide, almost completely microtrichose, except antepronotum and anterior margin of scutum. Postpronotal lobe with 1 seta and 2 groups of setulae: anteroventral group containing 7–9 rather stout and long setulae, and 6–7 appressed setulae around seta. Proepisternum with group of 6–7 long posterodorsally directed setulae on dorsal portion and 1 weak proepisternal seta on ventral portion. Prosternum consisting of very small subquadrate presternum and bare, microtrichose, transverse basisternum, which is twice as wide as high.

514 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Lateral cervical sclerite button-like, microtrichose. Mesonotum in profile slightly convex, scapular area abruptly bent. Scutum with numerous setulae not forming conspicuous rows except dorsocentral row; 8–10 setulae between dorsocentral lines. One acrostichal, 2 dorsocentral, 1 intra-alar, 1 postalar setae, 1 supra-alar seta midway between postalar seta and transverse suture. Spots of brown microtrichia at bases of setulae small and round; confluent spots forming larger spots anterior to suture (1–2 pairs, in P. corticalis often forming brown vittae), in area between branches of transverse suture, on postsutural portion along dorsocentral rows, and around prescutellar acrostichal seta. Scutellum conspicuously convex, with 2 pairs of scutellar setae and without any setulae, with pair of brown spots on disc and dots at bases of setae. Subscutellum in lateral view 0.33 times as long as scutellum. Subscutellum and mediotergite gray microtrichose, often with brown spots. Notopleural triangle mostly brown at transverse suture, usually with gray parts; 2 subequally long notopleural setae. Anepistenum gray microtrichose, with brown spots at bases of setulae; these spots more or less confluent, especially at ventral margin; anepimeron gray, at ventral margin brown. Katepisternum, meron, katatergite and anatergite almost completely brown. Numerous setulae of anepisternum rather dense and half as long as each of 2 anepisternal setae at posterior margin. Katepisternum with 1 seta at posterodorsal margin and 2–3 setae at ventral margin. Wing. Moderately broad. Tegula unmodified, with 6–7 setulae and 2–3 longer marginal setae. Costagial section of costa dorsoventrally widened, with dense irregular setulae and 1 ventroapical and 1 dorsoapical seta. Humeral section of costa with 5–6 irregular rows of moderately long and thin setulae. Section of costa between humeral break and apex of vein R2+3 with two rows of setulae: fine at anteroventral margin, thickened at anterodorsal margin to midway between apices of veins R1 and R2+3; remaining portions of costa microtrichose.

Subcostal vein at apex moderately bent towards anterior margin. Vein Rs bare. Vein R1 straight, bare on proximal half, with 18–24 setulae dorsally distal to subcosta apex. Vein R2+3 straight or inconspicuously sinuous. Vein R4+5 bare, ending at wing apex. Crossvein R-M almost perpendicular to vein R4+5, situated at level of middle of pterostigma. Vein M almost straight; its second (discal) section without stump vein posteriorly; its penultimate section of various lengths, being conspicuously variable in P. millepunctata, in which it varies from 1.2 to 2.7 times as long as crossvein R-M; distal section of vein M straight, not bent anteriorly. Crossvein

BM-Cu perpendicular to vein M. Crossvein DM-Cu straight. Vein Cu2 sharply bent outwards, its section between bend and vein A1 as long as or longer than section between fork of vein Cu and elbow; vein A1+Cu2 reaching posterior wing margin. Triangular posterodistal lobe of cell bcu shorter than apical section of vein A1+Cu2. Vein A2 well developed as fold. Alula moderately developed. Dorsal calypter with long marginal fringe, 1.5–2 times as long as ventral calypter. Halter unmodified, with rounded and short knob; knob surface with very short setulae. Legs. Neither thickened nor having unusual vestiture. Forecoxa with conspicuously flattened anterior surface and 4-5 long ventromarginal setae about twice as long as numerous setulae on anterior surface; foretrochanter unmodified, short and sparsely setulose; forefemur mostly dark, microtrichose, with sparse appressed setulae, 2 posterodorsal and one posteroventral rows of setae; basal 0.33 of foretibia brown, at apical third with dark ring. Midcoxa with row of 9–10 setae on eucoxa (anterior sclerite) and 2 setae on disticoxa (posterolateral sclerite); midcoxal prong well developed, rather stout and chitinized; midtrochanter rather elongate, moderately setulose, unmodified; midfemur short setulose, with anterior row of 6–10 short setae 0.4–0.5

515 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 6–11. Pseudotephritis, wings. 6. P. vau. 7. P. approximata. 8–9. P. millepunctata. 8. China: “Han- tao-he-tzu” (CM). 9. P. ussurica, type (from Krivosheina & Krivosheina, 1997c). 10–11. P. corticalis, syntypes, variations of wing pattern. times as long as width of femur and posteroventral row of 9–15 setae 0.8–0.9 times as long as width of femur; midtibia more or less darkened on basal half and with single dark ring on apical third; ventroapical seta twice as long as other thickened setae at apex of midtibia. Hindcoxa unmodified, with 9-12 setulae on eucoxa and 2–3 lateromarginal stronger setae, which are 1.5– 2 times as long as setulae. Hindtrochanter unmodified, setulose on ventral surface. Hindfemur unmodified, moderately setulose, with 2 subapical setae on dorsal surface. Hindtibia very slightly curved, otherwise unmodified, without erect setulae, setulae of ventroapical row short and very slightly thickened; basal half darkened without pale ring dividing it, apical third with wide dark ring. Tarsi of all legs without modifications; tarsomeres 1 and 2 of hindleg with yellowish setulae forming short brushes on posteroventral surfaces; otherwise setulae of tarsomeres more or less symmetrical. Claws simple; pulvilli small; empodium bare, shorter than longest setae of tarsomere 5. Abdomen. With broad, microtrichose, setulose tergites, well-developed microtrichose pleural membrane without any setulae and moderately narrow, microtrichose and setulose sternites. Tergites wider than long, with speckled brown-gray pattern; sternites 2–4 of male and 2–5 of female as long as wide, other preabdominal sternites wider than long. Well-developed spiracles situated on ventral aspect of abdomen, on pleural membrane close to tergite margins. Female

516 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) tergite 6 short and narrowed, microtrichose with brown spots and setulose, partially hidden underneath margin of tergite 5. Sternites 3–6 of female with very long and narrow anteromedial apodemes (Figs. 18, 19). Oviscape gray microtrichose or matt, with brown-gray pattern. Male terminalia. Segments 6-8 clearly microtrichose. Epandrium with long, slightly mesally curved surstylus with short mesal lobe, with 4–5 prensisetae and numerous setulae in P. vau, P. approximata, and P. millepunctata (Figs. 12, 13, 15), in P. corticalis surstylus short, with mesal lobe subequal to its basal portion, with 2–3 prensisetae. Proctiger large in P. vau, P. approximata, and P. millepunctata, and smaller in P. corticalis (Figs. 16, 17). Metaphallic plate narrow and long in P. vau, P. approximata, and P. millepunctata (Figs. 12–15), wide and short in P. corticalis (Fig. 17). Phallus long spinulose (Fig. 12). Female terminalia. Eversible membrane without visible scales. Aculeus long and narrow; cercal unit elongate oval, with long setulae. Three spherical spermathecae with short inverted necks (Figs. 20, 21). Included species and distribution (Appendix 3, Figs. 125–127) Eight available names have been proposed for species now assigned to this genus, as follows: Pseudotephritis vau (Say), 1830 (Ortalis) (Canada: Alberta; British Columbia; Northwest Territories (new record); Nova Scotia (new record); Quebec; USA: Alabama (new record); Arkansas; California; Connecticut (new record); D.C.; Florida (new record); Idaho; Indiana; Iowa (new record); Kansas (new record); Louisiana (new record); Maine; Maryland; Massachusetts; Michigan; Minnesota; Mississippi; Missouri; New Hampshire; New Jersey; New York; Ohio; Oklahoma (new record); Oregon (new record), Pennsylvania; Rhode Island; Texas; Virginia; Washington; West Virginia (new record)) (Fig. 125). P. corticalis (Loew), 1873 (Stictocephala) (Canada: Alberta; British Columbia (new record); Manitoba (new record); Ontario (new record); Quebec (new record); Saskatchewan (new record); USA: Connecticut; D.C.; Maryland; New Jersey; New York; Ohio; Pennsylvania; Utah (new record); Virginia; Norway; Russia: NW European Territory; N Caucasus; Far East (Figs. 126–127). P. approximata Banks, 1914 (as “P. appoximata”) (SE Canada, NE USA, Kansas (new record)) (Fig. 126). P. metzi Johnson, 1915 (NE USA). P. conjuncta Johnson, 1921 (as P. vau var. conjuncta) (NE USA). P. millepunctata (Hennig), 1939 (Myennis) (NE China (new record); E Russia) (Fig. 127). P. trypetoptera (Hennig), 1939 (Myennis) (NW and E Russia, Mongolia). P. ussurica Krivosheina and Krivosheina, 1997 (as “P. ussuruca” on p. 675, inadvertent error; as P. ussurica on pp. 677–678) (E Russia). P. vau var. californica Steyskal, 1962 and P. vau var. idahoana Steyskal, 1962 from western USA are unavailable names. The validity of some of these names needs further clarification. Johnson (1921) considered approximata, metzi and conjuncta as morphological varieties of P. vau. Malloch (1931) reassigned full species status to them and noted that these three species are close to each other, but not to P. vau. Steyskal (1961, 1965) contended that metzi and conjuncta are synonyms of P. vau, but left P. approximata as a valid species, based on its conspicuously larger size, having an additional dark spot in the costal cell, and a wider metaphallic plate. Indeed, there is a full range of intermediate forms of P. vau depending on the distance between crossveins R-M and DM-Cu and wing pattern intensity. We have no additional evidence either supporting the status of P. approximata as a valid species or for its identity with P. vau, and maintain the concept of

517 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 12–21. Pseudotephritis, terminalia. 12. P. approximata, male terminalia, anterior view (from Steyskal, 1962). 13. P. vau, epandrium, posterior view (proctiger removed). 14–15. P. millepunctata, epandrium, posterior view (proctiger removed) (modified from Krivosheina & Krivosheina, 1997c). 14. Specimen with approximated crossveins. 15. Type of P. ussurica. 16–17. P. corticalis, epandrium, posterior view. 16. Specimen from Far East Russia (modified from Krivosheina & Krivosheina, 1997c). 17. Specimen from northwestern USA. 18. P. vau, abdominal sternites 3–6 of female. 19. P. corticalis, abdomen, ventral view (oviscape removed). 20. P. vau, spermathecae. 21. P. corticalis, spermathecae.

518 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) this species proposed by Steyskal (1962). Hennig (1939) was unaware that the species he described as Myennis millepunctata and M. trypetoptera actually belong to a genus described from North America. Steyskal (1962) synonymized the nearctic P. corticalis (Loew, 1873) with the palaearctic P. trypetoptera (Hennig, 1939) and accompanied this action with the following comment: “Hennig (personal communication) confirms the above synonymy from comparison of American material sent him; his type was from Luga, Tolmatchevo, Russia (near Leningrad)”. This synonymy was accepted in both the nearctic (Steyskal, 1965) and palaearctic (Soós, 1984) catalogs. Later, Krivosheina and Krivosheina (1997c) resurrected the name P. trypetoptera (Hennig) based on the fact that “…Steyskal (1962) gives the only difference of P. corticalis (Loew) — surstylus with 2 clear denticles [= prensisetae] only. P. trypetoptera (Hennig) is characterized by a different type of surstylus morphology.” Indeed, the specimen figured by Krivosheina and Krivosheina (1997c: Fig. 1,4 and 1,7) differs from that figured by Steyskal (1962: Fig. 4). In fact, most nearctic specimens differ from palaearctic specimens in having a somewhat reduced wing pattern. Nevertheless, a study of a large series of North American specimens shows that both the wing pattern and number of prensisetae overlap: some specimens from the U.S.A. possess 3 prensisetae at least on one surstylus, as in specimens from Far East Russia (Krivosheina and Krivosheina, loc. cit. Fig. 1,7); the surstylus, proctiger and metaphallic plate shape differences between male terminalia figured by Steyskal and Krivosheina and Krivosheina are due to different angles and minor inaccuracies in drawing, and were found to be almost identical in our study. About 15% of the North American specimens have a wing patterns typical of palaearctic specimens (Fig. 11). We, therefore, again treat P. trypetoptera a synonym of P. corticalis. Finally, Krivosheina and Krivosheina (1997c) described P. ussurica based upon specimens differing from P. millepunctata in having more distantly spaced crossveins R-M and DM-Cu and widely separated dark spots lying on the crossveins. The material that we studied showed transitional forms between them (similar to series of variation in P. vau). The differences between forms of the epandrium resulted from different angles of view. Finally, the holotype of P. ussurica was from a series of typical P. millepunctata reared from under the bark of oak. Kameneva (2001) recently synonymized these two names. Determination. The four species recognized by us are included in the following key.

Key to species of Pseudotephritis

1. Wing tip hyaline from apex of vein R2+3 to apex of vein M except small dark spot at apex of

vein R4+5; dark area over crossvein R-M with hyaline spots on either side of crossvein (Fig. 10–11). Eye as long as high or longer than high (Fig. 5) ...... P. corticalis (Loew)

- Wing tip with complete brown band extending from apex of vein R2+3 to, or beyond, apex of vein M; dark area over crossvein R-M without hyaline spots (Figs. 6–9). Eye higher than long...... 2 2. Extension of cell bcu without dark brown spot (Fig. 9). Subbasal crossband dark brown, in cell dm narrower or, at most, as broad as in cell bm. Palaearctic species ...... P. millepunctata (Hennig) - Extension of cell bcu with dark brown spot which is conspicuously darker than subbasal crossband, which is yellow to yellow-brown (Figs. 6–7); width of latter in cell dm 1.5–2 times more than in cell bm. Nearctic species ...... 3

519 BIOTAXONOMY OF TEPHRITOIDEA

3. Costal cell with round brown spot between basal streak and larger dark subapical spot (Fig.

7). Apical crossband extending over vein R2+3 into cell r1. Wing length more than 5.2 mm...... P. approximata Banks - Costal cell between basal streak and larger subapical spot completely hyaline (Fig. 6).

Apical crossband at most touching apex of vein R2+3. Wing length less than 5.1 mm ...... P. vau (Say) Discussion The diagnostic characters of Pseudotephritis are either plesiomorphic or of uncertain polarity, although the presence of rather narrow and long metaphallic plate is a probable, but not proved synapomorphy of P. vau + P. approximata + P. millepunctata. The genus is rather heterogeneous because of the inclusion of P. corticalis that shares numerous characters with Callopistromyia. None of these characters is clearly a synapomorphy, but P. corticalis is a likely sister group of the whole Oedopa group of genera. We retain corticalis in Pseudotephritis (which is well-defined by plesiomorphies) provisionally, to avoid establishing one more monotypic genus based on uncertain relationships.

Callopistromyia Hendel, 1907 (Figs. 22-30, 128) Callopistria Loew, 1873: 62, 140. Type species: Platystoma annulipes Macquart, 1855 (by monotypy). Aldrich, 1905: 593; Williston, 1908: 277 (preoccupied name, non Callopistria Hübner, 1816, Lepidoptera). Callopistromyia Hendel, 1907: 98 (new name for Callopistria Loew); 1909a: 41; 1909b: 20; Curran, 1934b: 279; Aczél, 1951: 410; Steyskal, 1961: 405; 1965: 643; 1975: 147; 1979: 450; 1987: 807. The genus was established as monotypic and briefly diagnosed by Loew (1873, as Callopistria). Hendel (1909b) gave the most detailed description of this genus, based on the type species only. Steyskal (1975) transferred Pterocalla strigula Loew to Callopistromyia, which altered the concept of this genus, which has not been subsequently redescribed. Redescription Medium-sized (3.5–5.5 mm) flies with gray microtrichose body having speckled pattern of brown tomentum without shiny areas or bare dots at bases of setae (Fig. 23) and with setulose abdominal pleura (Fig. 28). Head. As wide as thorax, slightly higher than long (1.10–1.25; 1.00 in C. annulipes) or as high as long (C. strigula) and 1.4–1.6 times as wide as high. Lateral margins of frons converge anteriorly. Facial ridge with 4–6 setulae. Anterior part of gena with row of 6–8 dorsoclinate setulae reaching vibrissal edge and facial ridge. Gena 0.3–0.5 times as high as eye, densely setulose on posterior part. Eye slightly higher than long (1.1 : 1) (C. annulipes) or as high as long (C. strigula). One paravertical setula on each side of median occipital sclerite; 4-6 setulae in single postocular row; chaetotaxy otherwise similar to Pseudotephritis. Antennae well- separated, distance between their bases 2.0–2.2 times as wide as antennal socket. First flagellomere 1.0–1.1 times as long as wide. Proboscis and palpus similar to those of Pseudotephritis. Thorax. Generally as described for Pseudotephritis. Scutum with numerous irregularly

520 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) arranged setulae except more or less distinct dorsocentral row; 4–8 setulae between dorsocentral lines. Chaetotaxy similar to that of Pseudotephritis; 1 supra-alar seta closer to postalar seta than to transverse suture; intra-alar area without long setulae. Pattern of brown microtrichia as in Fig. 23. Subscutellum half as long as scutellum. Katepisternum almost completely brown, meron, katatergite and anatergite at least partially gray. Wings. Broad in C. annulipes (2.1 times as long as wide), narrow in C. strigula (2.8–3.0 times as long as wide). Crossvein R-M almost perpendicular to vein R4+5, situated slightly proximad

(in C. annulipes) or slightly distad (in C. strigula) of vein R1 apex. Vein M almost straight; its second section without stump vein; its penultimate section 1.7–3 times as long as crossvein R- M; ultimate section straight (Fig. 25) or slightly bent anteriorly (Fig. 24). Crossvein DM-Cu oblique, sinuous in C. annulipes, straight in C. strigula. Extension of vein Cu2 along vein A1

1.5–4.5 times as long as section between fork of vein Cu and elbow of vein Cu2. Posterodistal lobe of cell bcu triangular, longer than (in C. annulipes) or shorter than apical section of vein

A1+Cu2 (in C. strigula). Legs. Neither thickened, nor having unusual vestiture. All tibiae yellow at base and apex, with 3 dark rings. Midtibia with ventroapical seta more than twice as long as most other thickened setae at apex, and 2 shorter setae, about 0.7 and 0.6 times as long as longer seta, respectively. Tarsi of all legs without modifications. Abdomen. With well developed microtrichose pleural membrane bearing numerous setulae. Sternites 3–6 of female with very long and narrow anteromedial apodemes (Fig. 28). Oviscape gray microtrichose or matt, with brown gray pattern. Male terminalia. Surstylus short with mesal lobe almost equal to its basal portion, with 2–3 prensisetae; proctiger small; metaphallic plate moderately narrow, elongate (Figs. 26, 27). Phallus long spinulose. Female terminalia. Eversible membrane without scales. Aculeus long and narrow; cercal unit elongate oval, with long setulae (Fig. 29). Three spherical spermathecae with short inverted necks (Fig. 30). Included species and distribution (Appendix 3, Fig. 128) Callopistromyia annulipes (Macquart, 1855) (Canada: Alberta (new record); British Columbia; Ontario; United States: Alabama (new record); Arizona (new record); Connecticut; D.C.; Georgia; Idaho; Illinois; Indiana; Iowa (new record); Kansas; Kentucky; Louisiana (new record); Maine; Maryland; Massachusetts; Michigan; Mississippi; Montana; New Hampshire; New Jersey; New York; North Carolina; Ohio; Pennsylvania; Rhode Island; Tennessee; Utah; Vermont; Virginia; Washington; West Virginia; Wisconsin). C. strigula (Loew, 1873) (Canada: Manitoba, Ontario, Saskatchewan; United States: Colorado; D.C.; Columbia; Georgia; Illinois; Kansas; Maine; Maryland; Michigan; Minnesota; Nebraska; New Mexico; North Carolina; North Dakota; Pennsylvania (new record); South Dakota; Tennessee; Utah) (Fig. 128). Biology Steyskal (1979) collected puparia of C. annulipes and C. strigula in decaying cambium of a dead tree of boxelder (Acer negundo Linnaeus) and noted abundant adults of the first species on the bark of dying trees of Populus deltoides Barr. ex Marsh, ovipositing on dead trees of Robinia pseudacacia Linnaeus, and feeding on frass of wood borers in live trees of the latter

521 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 22–30. Callopistromyia. 22–24. C. annulipes. 22. Head, lateral view. 23. Mesonotum, dorsal view. 24. Wing. 25. C. strigula, wing. 26. C. annulipes, epandrium, posterior view. 27. C. strigula, epandrium, posterior view (from Steyskal, 1979). 28–30. C. annulipes. 28. Abdomen, ventral view (oviscape re- moved). 29. Female terminalia. 30. Spermatheca (1 of the 3).

522 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) species. Specimens from CNC have labels “dead poplar bark” on C. annulipes and “bleeding elm” and “wound on Acer negundo” on C. strigula specimens. Discussion The monophyly of the genus is supported by a single synapomorphy: the obtuse anterodistal angle of cell bcu. The two included species differ in wing shape and venation. Callopistromyia shares setulose abdominal pleura with other genera of the Oedopa group. It is the most generalized member of this group, and retains a speckled body pattern and head shape similar to Pseudotephritis corticalis, the possible sister-taxon to the entire Oedopa group. It is associated with dead wood of deciduous trees (similar to Pseudotephritis and genera of the Myennis group). Determination. Species can be determined with the following key.

Key to species of Callopistromyia

1. Wing less than 2.5 times as long as wide, oval. Crossvein R-M proximal of R1 apex (Fig. 24)...... C. annulipes (Macquart) - Wing more than 2.5 times as long as wide, with subparallel anterior and posterior margins.

Crossvein R-M distal of R1 apex (Fig. 25)...... C. strigula (Loew)

Oedopa Loew, 1873 (Figs. 31-42, 129) Oedopa Loew, 1868a: 287. Type species: Oedopa capito Loew, 1868 (by monotypy). Loew, 1873: 65, 146; Giglio-Tos, 1893: 12; 1895: 43; Cresson, 1906: 284; Hendel, 1910b: 63; Steyskal, 1961: 404; 1965: 653; 1974: 163; 1987: 806. Description Generic description has been given by Hendel (1910b: 63–65), to which we add the following.

Medium-sized (3.5–7.5 mm) flies with large head wider than long, bare vein R1, densely gray microtrichose body with sparsely microtrichose blackish areas or dots at bases of setae and setulose abdominal pleura. Head. Conspicuously wider than thorax, 1.5–1.7 times as high as long, wider than long, 3 times as wide as long. Frons wider than long, not protruding, in profile forming very obtuse angle with dorsal portion of face; lateral margins slightly convergent anteriorly. Frons white microtrichose, densely setulose, with small bare dots at bases of setulae to level of posterior ocelli, with unclear, narrow, white microtrichose median vitta broken by 3–4 bare spots. One lateroclinate and slightly reclinate ocellar seta and 3–5 smaller setulae on ocellar triangle. Two posterolaterally directed, approximated orbital setae; posterior seta slightly longer than anterior seta, additional mesoclinate seta often in line with anterior orbital seta. Frontal setulae all black, moderately stout, about 0.25–0.3 times as long as orbital setae. Face uniformly white microtrichose, slightly convex in profile in epistomal portion, without transverse furrow and with shallow and short antennal grooves. Clypeus low, with bulging, inverted horseshoe-shaped or tongue-like medial portion, and two sclerotized, flat lateral branches separated from it by folds, with paired swellings between them and peristomal margins. Parafacial very broad, 2–3 times as high as wide, with bare velvety black spot on dorsal 0.2, otherwise microtrichose, broadened on ventral half. Facial ridge with 2–3 setulae above vibrissal edge. Anterior part of

523 BIOTAXONOMY OF TEPHRITOIDEA gena with row of 4–8 lateroventrally directed setulae reaching ventral end of ptilinal suture, vibrissal edge and facial ridge. Gena 0.6–0.7 times as high as eye. Genal seta well developed. Eye as high as or very slightly higher than long, in live specimens horizontally banded. Vertex rounded, without ridge; postocellar seta divergent, slightly shorter than ocellar seta; medial vertical seta twice as long as postocellar seta; lateral vertical seta 0.7–0.9 times as long as medial seta. Occiput conspicuously concave dorsally, with very broad transverse brown or black mark, and slightly swollen ventrolaterally. No paravertical setula; 10-12 setulae in single postocular row; 3–4 long brown occipital setulae lateral to ventral portion of dorsal occipital suture and 3–4 setulae at level of occipital foramen. Anterodorsally curved supracervical setulae in two groups of 5-8 on each side separated by bare area. Postgena moderately white (in O. capito) or black (in O. ascriptiva) setulose. Hypostomal bridge (area between tentorial pits) twice as wide as high, bare, except lateroventral corners fine white setulose. Antennae well- separated, distance between their bases 2.5–2.9 times as wide as antennal socket. Scape very short. Pedicel wider than long, with inconspicuous medial lobe and dorsal incision; lateral surface with one row of marginal setulae, with 4 or 5 longest at ventral margin; 8-10 marginal setulae on medial surface forming 1–2 irregular rows; dorsal pedicellar seta almost as long as pedicel. First flagellomere 1.1–1.2 times as long as wide, short microtrichose. Two visible basal aristomeres. Terminal aristomere short pubescent. Proboscis stout, with rather small prementum 0.5 times as high as gena, bearing 4–6 setae on either side. Labellum fleshy. Palpus moderately narrow, sausage-shaped, curved dorsally, not broadened towards apex, with 6–8 setulae (as long as palpus width) at ventral margin and 5–8 shorter setulae on lateral and medial surfaces. Thorax. Robust, slightly longer than wide, microtrichose, except antepronotum and anterior margin of scutum, and subshiny dots at bases of setulae. Postpronotal lobe with 1 seta and 6–7 anteroventral setulae. Proepisternum with group of 6–10 fine long posterodorsally directed white and brown setulae on dorsal portion and 1 proepisternal seta on ventral portion subequal to dorsal setulae, but black and stouter. Prosternum consisting of very small presternum and bare, thickly microtrichose, trapezoidal basisternum, which is 1.5 times as wide as high. Lateral cervicalium microtrichose. Mesonotum in profile almost flat, scapular area abruptly curved. Scutum densely yellowish-white microtrichose, with 4 blackish, more sparsely microtrichose vittae and with numerous setulae forming one acrostichal row and one dorsocentral row on either side; 2–3 irregular rows of setulae between them; single supra-alar row well defined anterior and posterior to suture; between it and dorsocentral row 2–3 intra-alar rows of setulae separated by bare, microtrichose stripes from both dorsocentral and supra-alar row. One acrostichal, 1 dorsocentral, 1 intra-alar, 1 postalar and 1 supra-alar setae, latter inserted posterior to midpoint between postalar seta to transverse suture. Scutellum flattened, triangular, with 4 scutellar setae and without any setulae, with pair of large dark gray spots on disc and with thick microtrichia in middle and along margins. Subscutellum short, 0.2 times as long as scutellum. Subscutellum and mediotergite gray microtrichose. Notopleural triangle mostly dark gray, short and sparsely microtrichose at transverse suture, otherwise longer densely gray microtrichose; 2 subequal notopleural setae. Anepistenum gray microtrichose, with 2 dark gray longitudinal stripes of shorter microtrichia. Katepisternum microtrichose, sparser in middle; bases of setulae and setae with almost bare dots. Katepisternum with 1 seta at posterodorsal margin and 3–5 at ventral margin. Meron, katatergite and anatergite densely microtrichose, but at margins almost bare. Numerous setulae of anepisternum rather sparse and 0.33 times as long as 2 anepisternal setae at posterior margin.

524 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 31–42. Oedopa. 31–38. O. capito. 31. Habitus, dorsal view (from Loew, 1873). 32. Head, anterior view. 33. Habitus of syntype, lateral view. 34. Prothorax and forecoxae of syntype, anterior view. 35. Wing. 36. Male terminalia, posterior view. 37. Female preabdomen, ventral view (oviscape removed). 38. Spermathecae. 39–42. O. ascriptiva. 39. Head and mesonotum, dorsal view. 40. Head, anterior view. 41. Habitus, lateral view (from Hendel, 1910b). 42. Wing.

525 BIOTAXONOMY OF TEPHRITOIDEA

Wing. Tegula unmodified, with 4–5 setulae and 2–3 strong marginal setae. Costagial section of costa dorsoventrally widened, with rather sparse irregular setulae and 1 ventroapical seta. Humeral section of costa with 2–3 almost regular rows of moderately long and thin setulae.

Section of costa between humeral break and apex of vein R2+3 with two rows of subequal short and slightly thickened setulae to about apex of vein R2+3 (dorsal row slightly beyond apex of vein R2+3). Subcostal vein at apex slightly bent towards anterior margin. Vein R1 straight, bare.

Vein R2+3 slightly undulate. Vein R4+5 bare, ending at wing apex. Costal cell 2.8–3.0 times as long as pterostigma. Crossvein R-M almost perpendicular to vein R4+5, situated at level of middle of pterostigma. Vein M almost straight; its second section without stump vein; its penultimate section 5–6 times as long as crossvein R-M and 1.2–1.5 times as long as crossvein DM-Cu; ultimate section of vein M bent anteriorly. Crossvein BM-Cu perpendicular to vein M, clearly broken at anterior end. Crossvein DM-Cu sinuous, often with stump vein into cell dm.

Crossvein Cu2 with posterodistal lobe of cell bcu 2.2–2.7 times as long as section between fork of vein Cu and elbow of vein Cu2; vein A1+Cu2 reaching posterior wing margin. Posterodistal lobe of cell bcu triangular, as long as or slightly longer than apical section of vein A1+Cu2. Vein

A2 well developed as fold. Alula moderately developed. Dorsal calypter with long marginal fringe, 1.5–2 times as long as ventral calypter. Halter unmodified. Legs. Slightly thickened, white microtrichose. Forecoxa robust, with anterior surface flattened towards apex and 3-4 ventromarginal setae longer than setulae on anterior surface; foretrochanter unmodified; forefemur slightly swollen posterodorsally, and flattened at anteroventral surface, with slightly erect setulae and bare round spots at their bases, 2 posterodorsal and 1 posteroventral rows of setae; foretibia covered by short and appressed setulae, with brown ring at apex of basal third, another dark ring at base of apical third, both often inconspicuous in pale specimens. Midcoxa with row of 6–7 setae on elongate eucoxa and 3–4 setae on disticoxa; midcoxal prong well developed; midtrochanter unmodified; midfemur short setulose, with anterior row of 6–8 quite short setae 0.3–0.4 times as long as width of femur and with posterior row of 9–12 setae 0.5–0.6 times as long as width of femur; midtibia more or less darkened on basal half and with single dark ring on apical third; 2–3 ventroapical setae twice as long as other thickened setae at apex of midtibia, anterodorsal preapical seta developed and erect, but not much longer than surrounding setulae. Hindcoxa unmodified, with 4-5 anteroventral setae and 4–5 setulae on eucoxa and 2–3 stronger lateromarginal setae, 1.5–2 times as long as setulae. Hindtrochanter unmodified. Hindfemur without modifications, moderately setulose, with 2 subapical setae on dorsal surface. Hindtibia slightly curved, otherwise unmodified, with anterodorsal and posterodorsal rows of erect setulae, more conspicuous in O. ascriptiva, setulae of ventroapical row short and thickened; coloration like in foretibia and midtibia. Apical portion of foretibia, posteroventral surface of tarsomere 1 (and often 2) of foretarsus and anteroventral surface of tarsomeres 1–2 (–3) of hindtarsus with brushes of thick yellowish setulae; all tarsomeres with 2 rows of thickened setulae and 4 apical, elongate and thickened setulae on ventral surface, except 1 anteroventral row on foretarsomeres 1 and 2 and 1 posteroventral row on hindtarsomere 1; otherwise setulae of tarsomeres more or less symmetrical. Claws simple; pulvilli moderately developed; empodium bare or inconspicuously microtrichose, shorter than longest setae of tarsomere 5. Abdomen. Narrow oval, with broad, microtrichose, setulose tergites, well-developed microtrichose pleural membrane without any setulae, and moderately narrow, microtrichose

526 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) and setulose sternites. Tergites wider than long (except male tergite 5 almost as long as wide), with more or less distinct pattern of 4 rows of dark gray spots and dots at bases of setulae; sternites 3–4 of both male and female longer than wide, other sternites wider than long or subquadrate. Well-developed spiracles situated on ventral aspect of abdomen close to tergite margins. Female tergite 6 short and narrowed, microtrichose with brown spots and setulose, completely hidden underneath margin of tergite 5. Sternites 3–6 of female with very long and narrow anteromedial apodemes (Fig. 37). Oviscape gray microtrichose, with dots at bases of setulae. Male terminalia. Segments 6-8 microtrichose. Epandrium elongate oval, with large, slightly mesally curved surstylus, bearing 2 prensisetae and numerous setulae; proctiger short. Metaphallic plate large and long oval. Phallus short trichose (Fig. 36), without long spines. Female terminalia. Eversible membrane without visible scales. Aculeus long and narrow; cercal unit elongate oval, with moderately long setulae. Three mushroom-shaped spermathecae with inverted necks (Fig. 38). Included species and distribution (Appendix 3, Fig. 129) The genus includes 3 described species occurring in North and Central America: Oedopa capito Loew, 1868 (type-locality: Nebraska; Steyskal (1974) reported it from Arizona, California, Colorado, Idaho, Nebraska, New Mexico, South Carolina, Texas, Utah, Washington and Wyoming); O. ascriptiva Hendel, 1909 (Colorado) (Fig. 129) and O. elegans Giglio-Tos, 1893 (Mexico: Oaxaca). The latter species was originally briefly described in the key and redescribed in detail by Giglio-Tos (1895), but never redescribed or rediscovered since then. Its type specimens have not been located in MSNT (M. Daccordi, personal communication), and its placement in Oedopa remains provisional. Its redescription, with translation to English, is cited below. Oedopa ascriptiva is known with certainty only from Colorado, USA. The characters distinguishing it from O. capito given by Hendel (1909c) and Steyskal (1974) are not reliable, especially considering that both Hendel’s and Steyskal’s material from Colorado determined as O. capito contains darker specimens. These latter possess black and stout genal setulae as in typical O. ascriptiva (they are fine and short, yellow-brown in O. capito), but lack dark coloration of the scape and pedicel, and have dark spots in the pterostigma and posterior to it in cell r1, which is intermediate between most O. capito and O. ascriptiva. We assume that the form with stout black genal setulae belongs to O. ascriptiva, too, if the latter is not merely a morph of O. capito. Based on the examined material, the “typical” specimens of O. capito (with yellow genal setulae) are known from Canada: Alberta (new record) and USA: Arizona; California; Missouri (new record); Nebraska; Nevada (new record) and Washington. The form with black genal setulae and yellow pedicel and scape (O. sp. near ascriptiva) is known now from Canada: Alberta; British Columbia (new record of the genus); Saskatchewan (new record of the genus), and USA: Colorado and Utah. Other distributional records need to be rechecked; they are shown with white circles on the distribution map (Fig. 129). Biology Not known. The only association with biotopes that can be hypothesized from labels are prairies, dry places, or sand dunes in plains or low and middle altitude mountains. A label record “On Chilopsis” [shrub of the family Boraginaceae] apparently does not refer to a host plant.

527 BIOTAXONOMY OF TEPHRITOIDEA

Discussion The concept of this genus and its included species needs further revision. Monophyly of at least O. capito and O. ascriptiva is substantiated by the shape of the clypeus, which is unique in Ulidiidae, but it is not described for O. elegans. Relationships with the other genera of the Oedopa group have not yet been resolved. O. elegans is known from its very brief diagnosis and the more detailed redescription cited below. Its generic position, based on having a sinuous crossvein DM-Cu, is dubious, as the species has 2 black frontal spots as in Paroedopa punctigera. Determination Steyskal (1974) compiled the only complete key to species, which has been modified in the following key.

Key to species of Oedopa 1. Wing with numerous brown dots dispersed over whole disc. Frons with two round black spots ...... O. elegans Giglio-Tos - Wing with brown spots at anterior margin and apex only. Frons with black transverse bar ...... 2 2. Gena with inconspicuous yellow or brownish setulae ...... O. capito Loew - Gena with strong black setulae ...... 3 3. Pedicel and scape black ...... O. ascriptiva Hendel - Pedicel and scape yellow ...... O. sp. near ascriptiva Hendel Redescription of Oedopa elegans (after Giglio-Tos, 1895: 42, with translation into English) “… Oedopa elegans. Oedopa elegans Giglio-Tos (19), p. 12. Femmina. — Faccia cenerino-bruniccia; guancie larghe sparse di piccoli peli. — Proboscide nera; palpi bruno-giallicci. — Antenne gialle. — Fronte cenerino-bruniccia, leggermente ocracea in basso; due macchie nere rotonde laterali prima del vertice. — Torace bruno-cenerino, finissimamente punteggiato di bruno sul dorso. — Scudetto cenerino, appena ocraceo all’apice, munito di quattro setole, e con due macchie nere, piccole, rotonde nel mezzo. — Addome alquanto piatto, quasi conico, coi segmenti ben distinti, nero e tutto cosparso di pollinosità cenerina, con macchie irregolari, bruniccie; ovopositore largo, un po’ fulvo. — Piedi robusti, neri; le tibie giallo-fulve, colla base e due anelli l’uno verso il mezzo e l’altro presso l’apice, neri; tarsi gialli; uncini grandi, neri; ultimo articolo dei tarsi posteriori molto dilatato. — Ali ialine, macchiettate di moltissime macchie bruniccie, più intense verso la base; vene trasverse distanti, la posteriore bisinuosa. — Bilancieri gialli. — Lunghezza mm. 7. Una sola femmina. [Female. — Face grayish-brown; genae wide, with sparse small setulae. — Proboscis black; palpi brownish yellow. — Antennae yellow. — Frons grayish brown, slightly ochraceous at base; two black round spots laterally before vertex. — Thorax brownish gray, very finely brown dotted on dorsum. — Scutellum gray, very slightly ochraceous apically, bearing 4 setae, and with 2 black spots, in average small, round. — Abdomen quite flat, almost conical, with segments well distinct, black and completely gray microtrichose, with irregular brown spots; ovipositor wide, yellow. — Legs robust, black; tibiae creamy yellow, with two black rings, one

528 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) at middle, and the other at apex; tarsi yellowish; claws large, black; last joint of hindtarsi very dilated. — Wings hyaline, maculated with numerous brown spots, more intensively towards base; transverse vein distant, the posterior bisinuous. — Halter creamy. — Length 7 mm. One single female.] Hab. — Oaxaca (Sallé).”

Paroedopa Coquillett, 1900 (Figs. 43–48, 130) Paroedopa Coquillett, 1900: 22. Type species: Paroedopa punctigera Coquillett, 1900 (by original designation). Cresson, 1906: 284; Hendel, 1910b: 66; Curran, 1934b: 277; Steyskal, 1961: 404; 1965: 653; 1987: 806. Description See Hendel (1910b: 66). Principal differences from Oedopa are as follows. Head. Conspicuously wider than thorax, 1.7–1.8 times as high as long, 2.2–2.3 times as wide as long. Lateral margins of frons conspicuously divergent anteriorly. Frons matt, microtrichose only on posterior half. One reclinate ocellar seta and no setulae on ocellar triangle. Two reclinate orbital setae; posterior seta twice as long as anterior seta. Clypeus low, without separated anteromedial portion. Parafacial uniformly microtrichose. Anterior part of gena with row of 3–4 lateroventrally directed peristomal setulae and 2–3 setulae at vibrissal edge. Gena 0.25–0.3 times as high as eye. Eye 1.5 times as high as long. Vertex: postocellar seta slightly longer than ocellar seta. Occiput broadly darkened, microtrichose on dorsal portion, slightly produced posteriorly at eye margin and postgena, yellow on ventrolateral portion. No paravertical setula; 8–9 setulae in single postocular row; 0–1 long black occipital setulae lateral to ventral portion of suture. Supracervical setulae not examined. Postgena moderately black setulose. Antennae well-separated, distance between their bases twice as wide as antennal socket. Scape short. First flagellomere very short microtrichose. Terminal aristomere bare. Proboscis stout, with prementum as long as gena height. Palpus spoon-shaped, widened towards apex, with 4–5 setulae (conspicuously shorter than palpus width) at ventral margin and 4–5 at apex on lateral surface; 4–5 apicomedial setulae, twice as long as other setulae. Thorax. Coloration and vestiture similar to Oedopa. Postpronotal lobe with 1 seta and 3–4 anteroventral setulae. Proepisternum with group of 7–10 fine brown setulae on dorsal portion and 1 proepisternal seta on ventral portion, which is subequal to setulae, but stouter.

Wing. Vein R1 straight, bare. Vein R2+3 straight. Vein R4+5 ending slightly anterior to wing apex. Costal cell 1.5 times as long as pterostigma. Second section of vein M without stump vein; penultimate section of vein M 4–5 times as long as crossvein R-M and 1.5 times as long as crossvein DM-Cu. Crossvein BM-Cu oblique to vein M, constricted at anterior end. Crossvein

DM-Cu straight. Vein Cu2 with posterodistal lobe of cell bcu 3 times as long as section between fork of vein Cu and elbow of vein Cu2; vein A1+Cu2 reaching posterior wing margin.

Posterodistal lobe of cell bcu triangular, as long or slightly shorter than Vein A1+Cu2. Legs. Foretibia covered by short and appressed setulae and with posterodorsal row of longer erect setulae, with brown ring at apex of basal third, and another dark ring on apical third, both wide. Midcoxa with row of 6–7 setae on elongate eucoxa; disticoxa not examined; midcoxal

529 BIOTAXONOMY OF TEPHRITOIDEA prong well developed; midfemur as in Oedopa; midtibia banded like foretibia; 3–4 ventroapical setae twice as long as other 2–3 thickened setae at apex of midtibia and 1 anterodorsal preapical seta. Hindtibia slightly curved, otherwise unmodified, with anterodorsal and dorsal rows of erect setulae, setulae of ventro-apical row short and thickened; coloration like in fore and midtibia. Vestiture of tarsi like in Oedopa; apical 0.2 of tarsomere 1 and whole tarsomeres 2–5 dark brown. Abdomen. As in Oedopa. Male terminalia. Epandrium elongate oval, with large, mesally curved surstylus, bearing 2 long dentate prensisetae and numerous setulae; proctiger short. Metaphallic plate very large, long oval. Phallus trichose, without long spines. Female terminalia. Oviscape gray microtrichose, with dots around bases of setulae. Three mushroom-shaped spermathecae with inverted necks. Included species and distribution (Appendix 3, Fig. 130) Paroedopa punctigera Coquillett, 1900: 23 (USA: Arizona, New Mexico, Texas) (Fig. 130). Biology Unknown. Discussion Paroedopa and Oedopa share at least one synapomorphy, i.e., elongate abdominal sternite 4, plus several symplesiomorphies, which differentiate them from other Myennidini genera. Compared to Oedopa, the clypeus is unmodified (plesiomorphy) in Paroedopa, whereas the palpus is spoon-shaped and crossvein DM-Cu is straight (autapomorphies). The sister-group relationship between these genera is therefore quite probable, but see discussion of Stictomyia relationships below.

Stictoedopa Brèthes, 1926 (Figs. 49, 133) Stictoedopa Brèthes, 1926: 187. Type species: Stictoedopa ruizi Brèthes, 1926 (by monotypy). Description Known from the original French generic diagnosis and Latin description only, which are cited below (our English translation follows): “… Stictoedopa. Ce genre correspond presque aux genres Oedopa Loew et Paroedopa Coq., ayant avec le premier plus d’affinités qu’avec le second: les première et troisième veines longitudinales sont nues, la première cellule postérieure bien rétrécie vers le bord alaire, la cellule anale formant un angle aigu vers l’arrière, les joues avec un profil subcarré qui ont une hauteur presque égale à la hauteur verticale des yeux; il se rapproche de Paroedopa par les yeux plutôt elliptiques- verticaux, et par la veine transversale externe droite. Mais il diffère des deux genres cités par l’occiput légèrement surbaissé par rapport au niveax supérieur des yeux (comme dans les Rhopalomeridae), etc surtout par la trompe qui est filiforme, la partie basale pratiquement cachée dans l’ouverture buccale, avec les palpes presque rudimentaires à son extrémité et dont la partie externe est longue, un peu plus longue que la hauteur de la tête et terminée par deux

530 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs 43–49. Paroedopa and Stictoedopa. 43–48. P. punctigera. 43. Female, habitus, lateral view (from Hendel, 1910b). 44. Male, habitus, dorsal view. 45. Head, anterior view. 46. Wing. 47. Epandrium, posterior view. 48. Apex of surstylus, enlarged. 49. S. ruizi, wing (from Brèthes, 1926). labelles qui sont deux fois plus longues qu’épaisses. De plus, la veine médiastine atteint le bord costal après le milieu de l’aíle et sans que la veine costale apparaisse brisée à sa jonction. [This genus almost corresponds to genera Oedopa Loew and Paroedopa Coq., being more closely related to the first than to the second genus: the first and the third longitudinal veins are bare, the first posterior cell well narrowed towards wing margin, the anal cell forming an acute angle towards posterior [end], the cheeks in profile subquadrate, almost equal to vertical diameter of the eye; it is close to Paroedopa in the eye being rather vertical-elliptic, and in the external transverse vein being straight, but differs from these two genera in the occiput being slightly lowered compared to dorsal level of eyes (as in Rhopalomeridae), etc., but mostly in the proboscis filiform, its basal part practically hidden in the peristomal cavity, with the palpi almost rudimentary, to its [proboscis] apex whose external part is long, slightly longer than head height and ending in two lips that are twice as long as wide. Moreover, the mediastine vein reaches the costal margin after the

531 BIOTAXONOMY OF TEPHRITOIDEA middle of the wing and without appearing broken at junction.]”

“Stictoedopa Ruizi Brèthes, n. sp. Oculis (post mortem), triangulo ocellari et haustelle nigris; fronte testacea verticem versus rubescente; facie genisque albidis; antennis rubellis, pilis, seta et articule 3.0 apice nigris; thorace griseo, lineis 2 parallelis anterioribus et linea transversa alas versus nigris: scutello? Abdomine griseo, segmenta 2.0 macula antero-laterali et duabus ad marginem apicalem, segmenta 3.0 lineis duabus mediis apice extrorsum euntibus; segmenta 4.0 maculis 2 apice (et punctis nonnullis) nigris. Femoribus plus minus ferrugineis, uni-vel bi-annulato-fuscis; tibiis griseis biannulato-fuscis: tarsis plus minus ferrugineis, pilis nigris; unguibus longis, divaricatis, tenuibus, basi ferrugineis, apice nigris; pulvillis sut magnis, testaceis; alis subhyalinis, ad costam tantum fuscis, nigro sexplagiatis. Long.: 7 mm. Alae: 5 mm. [Eyes (post mortem), ocellar triangle and haustellum black; frons yellowish, towards vertex reddish; face and genae whitish; antennae pinkish, microtrichose, seta and apex of 3rd joint black; thorax gray, 2 parallel anterior lines and transverse line against wings black; scutellum? Abdomen gray, 2nd segment with anterolateral spot and two at apical margin, 3rd segment with double medial lines going towards external apices; 4th segment apex spot (and small dots) black. Femora more or less reddish, with one or two brown rings; tibiae with two brown rings: tarsi more or less reddish, setulae black; claws long, divergent, narrow, bases reddish, apices black; pulvilli large, yellowish; wings subhyaline, to costa brownish, with six black spots. Length: 7 mm. Wing: 5 mm.]” Included species and distribution (Appendix 3, Fig. 133) Stictoedopa ruizi Brèthes, 1926: 188 (Chile). Its type locality (“Hacienda ‘Las Mercedes’, près Camarico”) is not precisely located (each of three sites named Camarico in Chile are shown by a question mark in Fig. 133). Biology Unknown. Discussion Close to Paroedopa, differing mostly in details of venation (apical section of vein M curved anteriorly and ending before wing apex; triangular lobe of cell bcu twice as long as vein

A1+Cu2; and proboscis very narrow and long. This species is the only known representative of the Oedopa group in South America.

Stictomyia Bigot, 1885 (Figs. 50-59, 130) Stictomyia Bigot, 1885: clxvi [166]. Type species: Stictomyia longicornis Bigot, 1885 (by monotypy). Wulp, 1898: 377; Coquillett, 1900: 23; Aldrich, 1905: 594; Cresson, 1906: 284; Williston, 1908: 277; Hendel, 1910b: 60; Steyskal, 1961: 404; 1965: 654; 1987: 805. Description The generic description was given by Hendel (1910b: 61–63), and can be supplemented as follows. Medium-sized (4.0–5.0 mm) flies superficially resembling small beetles, with acute vertical ridge, concave occiput, round spotted wings, bare vein R1, slightly flattened and densely

532 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) microtrichose body with reticulate pattern of broadly confluent shiny black dots at bases of setulae, and setulose abdominal pleura. Head. Conspicuously wider than thorax, 1.5–1.8 times as high as long, 2–2.5 times as wide as long. Frons lateral margins slightly divergent anteriorly. Frons white microtrichose, densely setulose, with large bare dots at bases of setulae to level of posterior ocelli, median vitta nonsetulose. One reclinate and somewhat lateroclinate ocellar seta, without any additional setulae on ocellar triangle. Two posterolaterally directed orbital setae; posterior seta twice as long as anterior seta. Face convex in profile in epistomal portion, with velvety black medial spot between antennae and two shiny black swellings ventral to antennal bases, in S. longicornis with 2 callosities with violet sheen and transverse furrow below them (Figs. 50, 51). Clypeus moderately swollen, shiny black. Parafacial with shiny black callosity at frons border and bare matt black spot at middle (in S. longicornis) or with smaller shiny black spot on dorsal 0.2 and round shiny dots at setulae bases on ventral half (in S. punctata). Facial ridge with 2 setulae above vibrissal edge; 3 long peristomal setulae reaching ventral end of ptilinal suture. Gena 0.4–0.5 as high as eye. Genal seta well developed. Eye 1.4–1.5 times as high as long, in live specimens horizontally banded. Vertex with rather acute ridge; postocellar seta divergent, shorter than ocellar seta; medial vertical seta twice as long as postocellar; lateral vertical seta 0.8 as long as medial one. Occiput on dorsal portion deeply concave, wholly dark with microtrichia along eye margins only, and slightly swollen and yellow on ventral portion. Short paravertical setula between lateral vertical seta and suture, in line with vertical setae; 6–7 setulae in single postocular row. Other occipital setulae not examined. Postgena long but rather sparsely setulose, deeply excavated, gula wide and low. Antennae well-separated, distance between their bases twice as wide as antennal socket. Scape short in S. punctata, but as long as wide in S. longicornis; black setulose. Pedicel longer than wide, with inconspicuous medial lobe and dorsal incision in S. punctata, and with deep pedicellar notch in S. longicornis. First flagellomere three times as long as wide in S. punctata and 6 times longer in S. longicornis. Terminal aristomere bare. Proboscis with prementum almost as long as gena height, bearing 6–7 setae on each side. Palpus spoon-shaped apically, with 4–5 setulae (slightly shorter than average palpus width) at ventral margin and 4–6 longer subapical setulae on lateral and medial surfaces. Thorax. Robust, very slightly longer than wide, with reticulate pattern of microtrichia separated by shiny black confluent dots at bases of setulae and larger spots on antepronotum, postpronotum, notopleura and scutum. Postpronotal lobe with 1 seta and 7–10 setulae on anterolateral surface. Proepisternum with group of 4–7 long brown setulae on dorsal portion; proepisternal seta absent in S. longicornis; not examined in S. punctata. Scutum microtrichose, with pattern of partially confluent shiny black dots at bases of setulae; acrostichal and dorsocentral rows not differentiated from other setulae. 1–2 acrostichal, 1–2 dorsocentral, 1–2 intra-alar, 1 postalar, and 1 (rarely duplicated) supra-alar seta posterior to midpoint from postalar seta to transverse suture. Scutellum flat, rounded in dorsal view, subshiny black on disc, microtrichose in middle (in S. punctata) and along margins. Subscutellum very short, 0.2– 0.16 times as long as scutellum. Subscutellum and mediotergite sparsely microtrichose. Anepistenum gray microtrichose, without longitudinal bare stripes, but with isolated or partially fused bare dots at bases of setulae. Wing. Broad, with darkened disc and pattern consisting of darker spots at anterior margin similar to those in Oedopa and Paroedopa, and hyaline spots, smaller, more numerous and

533 BIOTAXONOMY OF TEPHRITOIDEA isolated in S. longicornis and larger, partially confluent and less numerous in S. punctata. Costagial section of costa with 3 rows of setulae and 1 ventroapical and often 1 dorsoapical seta 0.7–0.9 times as long as costagial section itself. Section of costa between humeral break and apex of vein R2+3 with two rows of thin setulae; remaining portions of costa microtrichose. Subcostal vein at apex sharply bent at obtuse angle towards anterior margin. Costal cell 1.4–2 times as long as pterostigma. Vein M with penultimate section 3–5 times as long as crossvein R- M and 1.7–1.9 times as long as crossvein DM-Cu; distal section of vein M bent anteriorly, almost as long as penultimate section in S. longicornis and 1.3 times as long as penultimate section in S. punctata. Crossvein DM-Cu slightly sinuous. Vein Cu2 with posterodistal lobe of cell bcu 1.5–3.0 times as long as section between fork of vein Cu and elbow of vein Cu2. Posterodistal lobe of cell bcu triangular, slightly shorter (in S. longicornis) or slightly longer (in

S. punctata) than Vein A1+Cu2. Alula well developed. Legs. Somewhat thickened, with white microtrichose pattern on coxae and femora. Forecoxa narrower and less flattened than in Oedopa and Paroedopa; 1 lateral preapical and 1–2 ventral preapical setae longer than 12–16 moderately long setulae on anterior surface; forefemur with slightly erect setulae and bare round spots at bases; foretibia apically widened, black with yellow ring beyond middle in S. longicornis or yellow with 3 dark rings, similar to Paroedopa, in S. punctata; preapical dorsal seta well developed in S. longicornis, not examined in S. punctata. Midcoxa with row of 6–7 stout setae on elongate eucoxa and 3–4 weaker setae on disticoxa; midfemur without rows of outstanding setae; midtibia yellow with 3 dark rings; 1 strong ventro-apical seta 1.5 times as long as 2 other long setae and twice as long as 3–4 shorter thickened setae at apex of midtibia; preapical dorsal seta well developed, equal to ventro-apical seta. Hindcoxa and hindtrochanter unmodified. Hindfemur without modifications, with 2 subapical setae on dorsal surface. Hindtibia like in Oedopa and Paroedopa, 3 dark rings well expressed. Short and thickened setulae of ventro-apical row (comb) whitish. Apicoventral portion of foretibia, posteroventral surface of tarsomere 1(-2) of foretarsus and anteroventral surface of tarsomeres 1–2 (–3) of hindtarsus with brushes of thick yellowish setulae blackish on distalmost tarsomeres; all tarsomeres with almost equal setulae, not forming conspicuous rows, and with 4 apical elongate and thickened setulae on ventral surface, except foretarsomere 1 with two apical setulae, midtarsomere 1 with two well-defined ventral rows of thickened setulae, and hindtarsomere 1 with one posteroventral row. Apices of tarsomeres 1 and 2, and almost entire tarsomeres 3–5 darkened. Abdomen. Wide oval, microtrichose with pattern of fused black spots on tergites, pleura, and sternites. Sternites 2–4 subquadrate, other sternites wider than long. Female tergite 6 almost hidden underneath margin of tergite 5. Sternites 5–6 of female with short anteromedial apodemes (Fig. 54). Male terminalia. Epandrium wide oval, with thick mesally curved surstylus, bearing 1 prensiseta in S. punctata and no prensisetae in S. longicornis; in the latter, mesal surface with numerous short thickened setulae; proctiger short. Metaphallic plate as long as, or slightly shorter than, proctiger, truncate in S. punctata and almost quadrangular, medially concave. Phallus trichose (Fig. 59). Female terminalia. Similar to those in Oedopa. Oviscape with silver microtrichia and large spots at bases of setulae. Three spherical spermathecae with long inverted necks (Fig. 55).

534 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 50–59. Stictomyia. 50–55. S. longicornis. 50. Habitus, lateral view (from Hendel, 1910 b). 51. Head, anterior view. 52. Male terminalia, ventral view. 53. Male terminalia, posterior view. 54. Female abdomen, ventral view. 55. Spermathecae. 56–59. S. punctata. 56. Habitus, dorsolateral view. 57. Head, holotype, anterior view. 58. Epandrium, posterior view. 59. Phallus.

535 BIOTAXONOMY OF TEPHRITOIDEA

Included species and distribution (Appendix 3, Fig. 130) Stictomyia longicornis Bigot, 1885: [166] (USA: California; Arizona; New Mexico; Texas; Mexico), S. punctata Coquillett, 1900: 23 (USA: Arizona (new record); Idaho (new record); New Mexico; Texas) (Fig. 130). Biology Hunter et al. (1913) reported of a rearing record of S. longicornis from rotting cacti. Discussion Despite its appearance, which deviates from the general morphological plan of the Myennidini, Stictomyia clearly belongs in this tribe and specifically in the Oedopa group of genera based on the following characters: presence of a metaphallic plate, setulae on the abdominal pleura, and more than one ventro-apical seta on the midtibia. Within the Myennidini, Stictomyia is probably the sister-group of Paroedopa: they both possess a preapical dorsal seta on the foretibia and a round black spot between the antennal bases, which are believed to be synapomorphies. Stictomyia differs from Paroedopa in having a subquadrate sternite 4, which is a plesiomorphic state in the Myennidini; if Stictomyia and Paroedopa are sister-groups, the broad sternite 4 in Stictomyia is hypothesized to be a reversal. The male terminalia are different from the Myennidini ground plan and even among the two included species; however, the monophyly of Stictomyia is well-supported by having an elongate first flagellomere, dark wing pattern with hyaline spots, and an apically truncate metaphallic plate. Determination The species can be determined using the following key.

Key to species of Stictomyia 1. First flagellomere less than 4 times as long as wide. Black spots ventral to antennal bases

flat. Cells r1, r2+3 and dm with 3–5 hyaline spots each (Fig. 56). Wing tip with hyaline spots. Male terminalia: one acute prensiseta developed; metaphallic plate truncate apically, but not bilobate (Fig. 58) ...... S. punctata Coquillett - First flagellomere more than 4 times as long as wide. Black spots ventral to antennal bases

on raised calluses. Cells r1, r2+3 and dm with series of 6-15 small hyaline spots each. Wing tip wholly dark (Fig. 50). Male terminalia: no prensisetae; metaphallic plate slightly bilobate at apex (Fig. 52–53)...... S. longicornis Bigot

Ulidiotites Steyskal, 1961 (Figs. 60-66, 129) Ulidiotites Steyskal, 1961: 406; Type species: Ulidiotites dakotana Steyskal, 1961 (by original designation). Steyskal, 1965: 649; 1987: 806. Redescription Rather small or medium-sized (wing length 2.8–4.2 mm, female body length 4.0–6.1 mm) flies with large rounded head, bare vein R1, gray, densely microtrichose body without sharp blackish areas or dots, and with setulose abdominal pleura. Steyskal (1961) gave an adequate original description, to which the following details can be added. Head. Lateral margins slightly divergent anteriorly. Frons matt, setulose, without small bare

536 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) dots. One short lateroclinate ocellar seta and no setulae on ocellar triangle. 1, rarely 2 posterolaterally directed orbital setae. Frontal setulae all black and thin, about 0.5 times as long as orbital setae. Face conspicuously protuberant anteriorly, matt and sparsely microtrichose, without transverse furrow and with shallow and short, but conspicuous antennal grooves. Clypeus low, receding and weakly sclerotized, without outstanding medial portion, with paired swellings between its posterolateral branches and margins of genae. Parafacial broad, with spots in posteroventral portion. Facial ridge with 4–6 setulae above vibrissal angle. Anterior part of gena with row of 4–6 lateroclinate setulae reaching ventral end of ptilinal suture, vibrissal edge and facial ridge. Gena 0.4–0.5 as high as eye. Genal seta barely differentiated from neighboring genal setulae. Eye very slightly higher than long, in dry specimens with slight pearly sheen. Vertex rounded, without ridge; postocellar seta divergent, almost twice as long as ocellars; both vertical setae twice as long as postocellar. Occiput on dorsal portion conspicuously concave, slightly swollen on lateroventral portion, without dark mark. No paravertical setula; 12-16 setulae in two postocular rows on each side. Supracervical setulae whitish. Postgena brown-black setulose. Antennae well-separated, distance between bases 2.2– 2.5 times as wide as antennal socket. Scape and pedicel short, with sparse and short brown to black setulae, dorsal seta twice as long as other setulae. First flagellomere 1.0–1.1 as long as wide, short microtrichose. Two visible basal aristomeres. Terminal aristomere almost bare. Proboscis short, with small prementum much shorter than gena height, bearing 4–6 setae on each side. Palpus narrow, not broadened towards apex, with 4–6 setulae (2–2.5 times as long as palpus width) on ventral margin and 2–4 at apex. Thorax. Robust, slightly longer than wide, uniformly microtrichose, except antepronotum. Postpronotal lobe with 1 seta and 5–6 anteroventral setulae. Proepisternum with group of 5–6 long brown setulae on dorsal portion and 1 proepisternal seta on ventral portion, which is 0.7 times as long as these setulae. Prosternum bare, thickly microtrichose. Lateral cervical sclerite microtrichose. Scutum densely yellowish-white microtrichose, often with 4–6 darker or sparser microtrichose vittae; setulae rather sparse, forming 1–2 acrostichal rows, and row of 2–4 presutural and 4–6 postsutural dorsocentral setulae on each side; single supra-alar rows of 4–5 setulae each anterior and posterior of suture; between supra-alar and dorsocentral row 1–2 irregular intra-alar rows of setulae. 1–2 acrostichal, 1 dorsocentral, 1 intra-alar, 1 postalar, all in line, close to scutellum, postsutural supra-alar seta not differentiated from setulae. Scutellum flattened, short triangular, with 4 scutellar setae, setulae lacking. Subscutellum comparatively large, 1/4 as long as scutellum. Two notopleural setae subequal in length. Pleura without dark stripes or bare dots, with sparse, black, moderately long setulae. Two anepisternal and 1 katepisternal seta. Wing. Tegula unmodified, with 2–3 setulae and 1–2 stronger marginal setae. Costagial section of costa with sparse irregular setulae and 1 ventro-apical seta. Humeral section with 2 almost regular rows of short thin setulae. Section of costa between subcostal break and apex of vein

R2+3 with two rows of setulae to apex of vein R2+3 (setulae of ventral row 1.5 times as long as setulae of dorsal row). Subcostal vein at apex slightly bent towards anterior margin. Vein R4+5 bare, ending anterior to wing apex. Costal cell twice as long as pterostigma. Crossvein R-M almost perpendicular to R4+5 vein, situated at middle of pterostigma. Vein M almost straight; its second section without stump vein; its penultimate section 4–6 times as long as crossvein R-M and 1.5–2.1 times as long as crossvein DM-Cu. Crossvein DM-Cu straight. Vein Cu2 with

537 BIOTAXONOMY OF TEPHRITOIDEA

posterodistal lobe of cell bcu as long as section between fork of vein Cu and elbow of vein Cu2; vein A1+Cu2 reaching posterior wing margin at least as fold. Posterodistal lobe of cell bcu triangular, 3–4 times shorter than apical section of vein A1+Cu2. Legs. Completely yellow without dark pattern, fine microtrichose, with long, sparse, erect and thickened setulae on tibiae, especially in female. Forecoxa with 1–2 ventromarginal setae longer than setulae on anterior surface; forefemur slightly swollen posterodorsally, with 5 rows of setae and long erect setulae almost equal in length; foretibia with rows of moderately sparse, slightly thickened, erect setulae. Midcoxa with row of 5–6 setae on eucoxa and 2–3 setae on disticoxa; midfemur short setulose, with anterior row of 3–4 setae and posterior row of 6–7 setae 0.4–0.6 times as long as width of femur, ventral surface bare; midtibia with 2 ventro- apical setae 1.3–1.5 times as long as other 4 thickened setae at apex of midtibia, anterodorsal preapical seta barely different from other erect setulae of dorsal rows. Hindcoxa unmodified, with 8–10 long setulae and almost equal setae. Hindfemur conspicuously swollen dorsally, long and sparsely setulose, with 3 subapical setae on dorsal surface and row of 4–5 anteroapical setae longer than width of femur. Hindtibia slightly curved, with anterior and anterodorsal rows of thickened erect setulae, 0.6–0.8 times as long as tibia width and 2-3 thickened apical anterodorsal setae almost as long as tibial width. Brushes of thick yellowish setulae present on posteroventral surface of foretibia (apical portion), tarsomere 1 of foretarsus, and anteroventral surface of tarsomeres 1–2 of hindtarsus; tarsomeres 1–3 of all legs with erect long setulae on dorsal surface; midtarsomere 1 with thickened setulae forming 4–5 rows on ventral surface, tarsomeres 2–4 with 2–4 rows of much shorter but also thickened setulae on ventral surface, 1 anteroventral row on foretarsomeres 1 and 2, and 1 posteroventral row on hindtarsomere 1. Abdomen. Narrow oval, with broad microtrichose, setulose tergites, well-developed microtrichose pleural membrane with numerous long setulae and moderately narrow, microtrichose and setulose sternites. Female tergite 6 exposed and moderately large (Figs. 64– 65). Sternite 4 of female longer than wide, other preabdominal sternites wider than long or subquadrate; sternites 4–6 of female with anteromedial apodemes (Fig. 66). Male terminalia. Segments 6–8 microtrichose. Epandrium elongate oval, with slightly mesally curved surstylus, bearing numerous long and rather thick setulae with prensisetae barely differentiated among them; proctiger short. Metaphallic plate short and small. Phallus long trichose (Fig. 63). Female terminalia. Oviscape microtrichose; aculeus long and narrow; cercal unit laterally flattened, acute in dorsal view, with long setulae. Three round spermathecae. Included species and distribution (Appendix 3, Fig. 129) Ulidiotites dakotana Steyskal, 1961 (USA: Kansas (new record); Mississippi (new record); Nebraska; Nevada (new record); South Dakota; Utah (new record); Wyoming (new record) (Fig. 129). Biology Unknown. Discussion This genus possesses both setulose abdominal pleura and more than one apicoventral thickened setae on the midtibia. It belongs to the lineage within the Oedopa group, which also

538 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 60–73. Ulidiotites and Namibotites. 60–66. U. dakotana. 60. Habitus, lateral view. 61. Head and mesonotum, dorsal view. 62. Head, anterior view. 63. Male (Utah), terminalia, posterior view. 64. Female (Utah), abdomen, dorsal view. 65. Female (Nebraska), dissected abdomen (aculeus obliquely com- pressed). 66. Same, sternites 3–6, ventral view. 67–73. N. argentata (from Barraclough, 2000). 67–68. Head. 67. Lateral view. 68. Anterior view. 69. Wing. 70–71. Male terminalia. 70. Left lateral view. 71. Posterior view. 72. Apex of aculeus. 73. Spermatheca (1 of 2).

539 BIOTAXONOMY OF TEPHRITOIDEA includes Oedopa, Paroedopa, Stictomyia and, very probably, Stictoedopa. All five genera share the following two characters: vein R1 bare, and gena high and setulose. Ulidiotites, Oedopa and Paroedopa also share the elongate oval epandrium. Along with some autapomorphies (body and wing pattern absent, posterodistal lobe of cell bcu short, body vestiture sparse, long and fine), Ulidiotites differs from the other genera also by its long trichose epandrium and large female tergite 6; both characters believed to be plesiomorphic with certain probability of reversal. We therefore consider Ulidiotites a sister-group of the Oedopa + Paroedopa + Stictomyia lineage, but this must be confirmed by additional characters. Newly discovered material has expanded the known distribution of this genus. At the same time, the considerable morphological variation within the new material makes further detailed study necessary. Specimens with 2 and 4 rows of acrostichal setae and more and less erect tibial setulae apparently belong to different size classes or sexes of the same species, but a thorough examination of genital characters is needed to confirm this.

Namibotites Barraclough, 2000 (Figs. 67–73) Namibotites Barraclough, 2000: 94. Type species: Namibotites argentata Barraclough, 2000 (by original designation). Description See Barraclough (2000). Included species and distribution Namibotites argentata Barraclough, 2000: 94 (Namibia). Discussion Namibotites shows strong superficial similarity to Ulidiotites in its uniformly microtrichose body and hyaline wing with bare vein R1 and vein M bent anteriorly. Otherwise it does not have diagnostic characters of the Oedopa group: the midtibia has one long ventroapical seta (although 3 short setae also present), and the abdominal pleura have no setulae (Barraclough, personal communication). The disjunct ranges of Namibotites and Ulidiotites (in the arid areas of Africa and North America, respectively), which is unusual for closely related tephritoids, also suggests remote relationships. Many characters of Namibotites are autapomorphic, and its phylogenetic relationships within the Myennidini are uncertain.

Arborotites Barraclough, 2000 (Figs. 74–82) Undescribed genus: Barraclough, 1995: 129. Arborotites Barraclough, 2000: 85. Type species: Arborotites stuckenbergi Barraclough, 2000 (by original designation). Description See Barraclough (2000). Included species and distribution Arborotites stuckenbergi Barraclough, 2000 (South Africa: Mpumalanga, KwaZulu–Natal, East Cape; Zimbabwe: North Vumba in eastern highlands).

540 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 74–82. Arborotites stuckenbergi (from Barraclough, 2000). 74–75. Head. 74. Lateral view. 75. Anterior view. 76. Mesonotum, dorsal view. 77. Wing. 78. Femur. 79. Male terminalia, left lateral view. 80. Epandrium, posterior view. 81. Apex of aculeus. 82. Spermatheca (1 of 3).

541 BIOTAXONOMY OF TEPHRITOIDEA

Biology Adults were observed on shaded tree trunks in woodland, where the dominant tree species were Ozaroa obovata, Celtis africana and Dombeya burgessica (Barraclough, 2000). Discussion Arborotites retains plesiomorphic characters of Pseudotephritis: microtrichose speckled appearance and wing pattern, and absence of shiny areas on mesonotum and abdomen. It differs by the presence of a cluster of long intra-alar setulae, which is considered a synapomorphy with the Myennis + Pseudotephritina lineage, as well as by characters of unresolved polarity (long extension of cell bcu) and by autapomorphies (proximal position of crossvein DM-Cu, brown medial discal spot on scutellum, yellow subapical ring on femora). The short and wide metaphallic plate is apparently a plesiomorphy shared with Pseudotephritis corticalis, but not with P. vau, P. approximata and P. millepunctata.

Myennis Robineau-Desvoidy, 1830 (Figs. 83–96, 131–132) Myennis Robineau-Desvoidy, 1830: 717. Type species: Myennis fasciata Robineau-Desvoidy, 1830 (= Musca octopunctata Coquebert, 1798) (by monotypy). Loew, 1868b: 8; Rondani, 1869: 37; Becker, 1905: 104; Hendel, 1909a: 10; 1909b: 5, 11; Aczél, 1951: 403; Hering, 1956: 87; Mamaev et al., 1977: 245; Soós, 1984: 58; Krivosheina and Krivosheina, 1997b: 628; Krivosheina, 1998: 95; Kameneva, 2001: 159. Miennis: Rondani, 1869: 28 (unjustified emendation or error).

Both detailed redescriptions of Myennis compiled by Hendel (1909b: 11) and Hennig (1939: 72) refer to a much wider concept of the genus, which included, in the first case, species now referred to Neomyennis, and, in the second case, some species now in Pseudotephritis. Krivosheina and Krivosheina (1997b) provided no redescription of the genus, and we redescribe it below. Redescription Medium-sized (4.2–6.0 mm) flies similar to Pseudotephritis in head and body shape and in wing venation, with gray microtrichose body and shiny scutellum, but no speckled pattern. Head. As wide as thorax, 1.2–1.3 times as high as long and wider than high. Eyes 1.3–1.6 times as high as long. Frons in profile forming obtuse angle with dorsal portion of face; lateral margins parallel. Frons in M. octopunctata and allied species white microtrichose, with brown spots and small bare dots at bases of setulae; posterior third of frons uniformly microtrichose, with clearly expressed non-setulose median triangle narrowed into stripe reaching anterior margin; in M. sibirica Portschinsky frontal vitta matt, with sparse brown microtrichia and somewhat denser white microtrichia at eye margin. One lateroclinate and slightly proclinate ocellar seta. Two orbital setae, posterior longer and thicker than anterior. Frontal setulae black, 0.25–0.50 times as long as orbital setae. Face as in Pseudotephritis. Clypeus convex, about as high as first flagellomere width, horse-shoe shaped, in profile as long as palpus and 2-5 times as long as high, completely microtrichose or, in M. sibirica, with bare horizontal stripe. Parafacial with bare spot on dorsal 0.2, otherwise microtrichose. ventral portion of facial ridge (vibrissal angle) with 2–6 setulae. Anterior part of gena with row of 5–8 dorsoclinate setulae reaching

542 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 83–96. Myennis. 83–86. M. octopunctata. 83. Male, habitus (from Coquebert, 1798). 84. Wing. 85. Epandrium, posterior view. 86. Spermathecae. 87. M. tricolor, syntype male, habitus (from Hendel, 1909b). 88. M. mandschurica, wing (from Hering, 1956). 89–96. M. sibirica. 89. Head, lateral view. 90. Wing. 91. Epandrium, posterior view. 92. Male terminalia, lateroventral view. 93. Ejaculatory apodeme. 94. Female sternite 6. 95. Apex of aculeus. 96. Spermathecae.

543 BIOTAXONOMY OF TEPHRITOIDEA vibrissal edge and facial ridge. Gena 0.25–0.45 times as high as eye. Genal seta well developed. Eye higher than long. Vertex rounded, without ridge, microtrichose in M. octopunctata and allied species, or shiny in M. sibirica; postocellar seta divergent, as long as ocellars; vertical setae as in Pseudotephritis. Occiput white microtrichose or bare, otherwise as in Pseudotephritis, without transverse dark mark or dots at bases of postocular and occipital setulae. One paravertical setula on each side of median occipital sclerite or none; 7–10 setulae in single postocular row. Anterodorsally curved supracervical setulae in two groups of 14–18 on each side separated by bare area. Postgena moderately long setulose. Hypostomal bridge short and bare. Antennae: distance between bases 1.2–1.7 times as wide as antennal socket. Scape short. Pedicel wider than long, similar to that in Pseudotephritis. First flagellomere 1.1– 1.5 times as long as wide, short microtrichose. Two visible basal aristomeres. Terminal aristomere short pubescent. Proboscis stout, with large, rounded prementum that bears 10–12 setae on each side. Labellum fleshy, large. Palpus wide towards apex, with 6-7 setulae at ventral margin and 5–10 setulae on lateral and medial surfaces, all shorter than palpus width, and 2–3 ventral setae almost as long as palpus width. Thorax. Longer than wide, with gray and brown microtrichose areas, except antepronotum, anterior margin of scutum, supra-alar areas posterior of wing base, most of scutellum, subscutellum and mediotergite. Postpronotal lobe with 1 seta and 8–12 setulae. Proepisternum with group of 6–12 long posterodorsally directed setulae on dorsal portion and 1 weak proepisternal seta, slightly shorter than these setulae, on ventral portion. Prosternum as in Pseudotephritis. Scutum with conspicuous dorsocentral row of setulae, 8–10 setulae forming irregular rows between dorsocentral lines and 3–5 irregular rows between supra-alar and dorsocentral lines. One acrostichal, 2 dorsocentral, 1 intra-alar, 1 postalar setae and 1 postsutural supra-alar seta midway between postalar seta and transverse suture. Female with long setulae between postalar and dorsocentral setae, and intra-alar seta reduced, hardly different from surrounding setulae; in male, intra-alar seta as long as dorsocentrals and setulae as long as on other parts of mesonotum. Bases of setulae without small spots of brown microtrichia; large brown areas at transverse suture, and mesal and lateral to dorsocentral row, sometimes at middle of transverse suture; in M. sibirica, mesonotum mostly with brown microtrichia. Scutellum convex, with 4 scutellar setae (basal slightly shorter than apical) and without any setulae, bare, slightly shiny, with triangular anterior area and medial microtrichose vitta. Subscutellum 0.33 times as long as scutellum. Notopleural triangle sparsely brownish microtrichose in M. sibirica or uniformly gray microtrichose in M. octopunctata and allied species. Anepistenum gray microtrichose, with sparse spots at bases of setulae in M. octopunctata and allied species or sparsely brown microtrichose in M. sibirica; anepimeron, katepisternum, meron, katatergite and anatergite microtrichose similarly to anepisternum.

Wing. Venation as in Pseudotephritis, except crossvein R-M at or distal to vein R1 apex. Vein

R1 straight, bare on proximal half, distal to subcosta apex with 15–27 setulae on dorsal surface.

Crossvein R-M oblique to vein R4+5. Penultimate section of vein M shorter than or, as long as,

R-M. Crossvein DM-Cu straight, oblique or bent towards apex, but not sinuous. Vein Cu2 closing cell bcu, sharply bent, with posterodistal lobe of cell bcu as long as (in M. sibirica), or longer than, section between fork of vein Cu and elbow of vein Cu2 (in other species).

Posterodistal lobe of cell bcu triangular, shorter than apical section of vein A1+Cu2. Calypteres and halter as in Pseudotephritis.

544 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Legs. As described for Pseudotephritis, except femora non-microtrichose in M. sibirica, and tibiae uniformly yellow or brownish yellow without dark rings. Abdomen. With tergites at least partially setulose and microtrichose; pleural membrane microtrichose, without any setulae; sternites moderately narrow, microtrichose and setulose. Tergite 1 broadly shiny, tergite 4 with lateral shiny spots; other tergites broadly gray or brown microtrichose; in M. sibirica posterior third shiny; disk of female tergite 5 almost devoid of setulae, only marginal setae developed. Female tergite 6 short and narrowed, hidden beneath margin of tergite 5. Sternites 4–6 of female with long anteromedial apodemes (Fig. 94). Male terminalia. Segments 6–8 very sparsely microtrichose. Epandrium with slightly mesally curved surstylus with 3 prensisetae and numerous setulae, and with apical lobe (distal to prensisetae) almost as long as basal portion of surstylus. Proctiger short (Fig. 92). Metaphallic plate moderately developed, rounded at apex (Fig. 85, 91). Phallus long spinulose. Female terminalia. Oviscape sparsely microtrichose or matt. Eversible membrane without visible scales. Aculeus long and narrow; cercal unit elongate oval, with long setulae. Three almost spherical spermathecae with short inverted necks (Fig. 86, 96). Included species and distribution (Appendix 3, Fig. 131–132) Four species: Myennis octopunctata (Coquebert, 1798) (Musca) (= Scatophaga fasciata Fabricius, 1805) (Europe, except north; Turkey; Armenia (new record); Israel (new record); northern Kazakhstan; southern West Siberia; southern Far East Russia) (Figs 131–132); M. sibirica Portschinsky, 1892 (southern Far East Russia); M. tricolor Hendel, 1909 (= M. nebulosa Krivosheina and Krivosheina, n. syn.) (Kyrghyzia; Tajikistan; Turkmenistan; Uzbekistan); M. mandschurica Hering, 1956 (Russia: southern Far East; China: Heilongjiang) (Fig. 132). The record of M. octopunctata from the Franz-Josef Land (Soós, 1957) is obviously based on specimens occasionally introduced with poplar logs. The name “myennis Coquebert, 1798” appeared in the systematic database of Musca species names (Thompson and Pont, 1994: 99) without a reference to the page of publication or to the description of M. octopunctata, and not combined with any genus name (indirectly, it must mean Musca myennis, but such a binomen never appeared, even in the latter paper). The original paper (Coquebert, 1798) does not mention the name “myennis” either in the text or in the figure caption. According to Thompson and Pont (personal communication), the name “myennis” was included in the database as a specific epithet erroneously; it does not meet requirements of availability of the Code and is therefore an unavailable name. The description of M. nebulosa was based upon darker specimens of M. tricolor, having an intensive presutural vitta on the mesonotum, abdominal tergite 1 black and the subbasal crossband of the wing ending before vein A1+Cu2; no additional differences were found. A series of specimens from Samarkand has intermediate combination of character states: well- developed presutural vitta, yellow tergite 1, and subbasal crossband not extending beyond vein

A1+Cu2, or weak presutural vitta, dark brown tergite 1, and subbasal crossband extending well beyond vein A1+Cu2. These species cannot be distinguished and we therefore consider them synonyms. The status of M. mandschurica remains debatable. It is probable that both M. mandschurica and M. tricolor are either vicariant species or subspecies of M. octopunctata. However, the discovery of M. octopunctata in Far East Russia and specimens with intermediate wing pattern

545 BIOTAXONOMY OF TEPHRITOIDEA characters (Krivosheina, 1998) demonstrates the sympatric coexistence of M. octopunctata and M. mandschurica there, possibly with hybridization. M. octopunctata is believed to be occasionally introduced there from Europe, like some other synanthropic tephritoids. However, no conclusion can be made without the study of additional material. Hendel (1909a, 1909b) also assigned the following species to Myennis: M. appendiculata Hendel, 1909, M. zebra Hendel, 1909 (Brazil), M. cyaneiventris Hendel, 1909 currently placed in the genus Neomyennis Hendel, and M. nigra Hendel, 1909 (Peru; Costa Rica), recently transferred to the genus Terpnomyennis Kameneva (Pterocallini) (Kameneva, 2004), and M. scutellaris (Wiedemann, 1830), the type species of the genus Pseudodyscrasis Hernández. Biology Myennis octopunctata is associated with poplars in Europe and Israel. Vos-de-Vilde (1935) collected larvae of this species under poplar bark in the Netherlands. N.P. Krivosheina (Mamaev et al., 1977: 247) reared M. tricolor from larvae living under the bark of dead turanga (Populus euphratica Olivier) in Turkmenistan (Deynau near Chärjew). Determination Species of Myennis can be determined by the following key.

Key to species of Myennis 1. Wing with brown base and almost completely hyaline cell bcu, without crossband reaching

vein A1+Cu2, but with other 4 dark brown crossbands; preapical crossband (between apical

portion of cell r1 and middle of cell m) well developed (Fig. 90). Thorax and abdomen brown microtrichose with large shiny areas...... M. sibirica Portschinsky - Wing with yellow base and yellow, sometimes faint, subbasal crossband reaching cell bm to

middle of vein A1+Cu2 and 3 yellow crossbands: preapical crossband reduced, at most

represented by yellowish area at apex of cell r1, either separate or completely fused with apical band (Fig. 83, 84, 87, 88). Thorax and abdomen black to yellow, gray microtrichose with brown microtrichose spots and bare, shiny areas...... 2 2. Apical wing band much paler than other crossbands, with diffuse margins, yellow area at

apex of cell r1 fused with apical band (Fig. 87)...... M. tricolor Hendel - Apical band as dark as other crossbands on wing, with sharp margins, yellow area at apex of

cell r1 either broken into 2 spots or fused with apical band...... 3

3. Apical band entire: yellow areas on cell r1 apex and at wing apex fused (Fig. 88)...... M. mandschurica Hering

- Apical band broken into 2 spots: yellow areas on cell r1 apex and at wing apex separated (Fig. 83, 84)...... M. octopunctata (Coquebert)

Pseudotephritina Malloch, 1931 (Figs. 97-105, 134) Pseudotephritina Malloch, 1931: 5. Type species: Stictocephala cribellum Loew (by original designation). Aczél, 1951: 412; Steyskal, 1962: 257, 261; 1965: 647 (as subgenus of Pseudotephritis Johnson); 1975: 148; 1987: 807 (full generic rank). Redescription Medium-sized (5.0–7.5 mm) flies resembling Pseudotephritis in having gray microtrichose

546 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) body and non-setulose abdominal pleura, but having mesonotal pattern of bare brown-black dots at bases of setae, 1 orbital seta, vein M with spur vein into cell dm, and scutellum with 2 shiny spots. Only a brief original diagnosis has ever been published (Malloch, 1931), and the genus is redescribed below. Head. As wide as thorax, 1.3 times as high as long and 1.5 times as wide as high. Eye 1.1–1.2 times as high as long. Frons as in Myennis, microtrichose, with brown bare dots at bases of setulae; posterior third of frons uniformly microtrichose, with clearly expressed median microtrichose triangle narrowed into stripe reaching middle of frons. Ocellar setae as in Myennis. One orbital seta. Face as in Pseudotephritis. Clypeus in profile about 0.3 times as high as width of first flagellomere, completely microtrichose. Parafacial with bare spot on dorsal 0.16, otherwise microtrichose. Ventral portion of facial ridge (vibrissal angle) with 0–1 setula. Anterior part of gena with row of 4–6 inconspicuously dorsoclinate, rather short setulae reaching vibrissal edge. Gena 0.25–0.30 times as high as eye. Genal seta moderately developed. Eye 1.1 times as high as long. Vertex rounded, without ridge, microtrichose; postocellar seta divergent or parallel, as long as ocellars; vertical setae as in Pseudotephritis. Occiput white microtrichose, as in Pseudotephritis, without transverse dark mark but with brown dots at bases of postocular setulae. One paravertical setula or none; 5–7 setulae in single postocular row. Other occipital and supracervical setulae not examined. Postgena short and sparsely setulose. Hypostomal bridge short and bare. Antenna similar to that in Myennis and Pseudotephritis; distance between antennal bases 1.7–2.0 times as wide as antennal socket. First flagellomere 1.3–1.5 times as long as wide. Proboscis and palpus as in Myennis. Thorax. Longer than wide, mostly gray microtrichose, except antepronotum, anterior margin of scutum, two spots of scutellum, lateral portions of subscutellum and ventral margin of mediotergite shiny; numerous shiny black or brown dots at bases of setulae. Postpronotal lobe with 1 seta and 8–12 setulae. Proepisternum with group of 12–18 long and fine, posterodorsally directed setulae on dorsal portion and 1 very weak proepisternal seta, which is 0.33 times as long as these setulae, on ventral portion. Prosternum and chaetotaxy of scutum as described for Pseudotephritis and Myennis. Female, instead of intra-alar seta, with series of 4–6 setulae 0.5– 0.6 as long as dorsocentral seta; male with intra-alar seta as long as dorsocentrals. Posterior intra-alar corner of scutum and lateroventral surface of scutellum covered by thick and short velvety-black microtrichia. Brown areas both laterally and at middle of transverse suture, and mesal and lateral to dorsocentral row, sometimes also on supra-alar area. Scutellum convex, with 2 pairs of scutellar setae (basal seta conspicuously shorter than apical seta and almost aligned with apical seta and dorsocentral seta), bare, almost shiny, with microtrichose triangular anterior area and medial vitta. Subscutellum 0.33 times as long as scutellum. Notopleural triangle as in Pseudotephritis. Anepistenum and katepisternum gray microtrichose, with isolated brown dots at bases of setulae; anepimeron and meron, katatergite and anatergite uniformly gray microtrichose, sometimes with few brown spots.

Wing. Venation as in Pseudotephritis. Vein R1 straight, bare on proximal half, distal of subcosta apex with 20–30 setulae on dorsal surface. Penultimate section of vein M as long as crossvein R-M. Crossvein DM-Cu not sinuous. Vein Cu2 with posterodistal lobe of cell bcu longer than section between fork of vein Cu and elbow of vein Cu2. Posterodistal lobe of cell bcu shorter than A1+Cu2. Calypteres and halter as in Pseudotephritis.

547 BIOTAXONOMY OF TEPHRITOIDEA

Legs. As described for Pseudotephritis, except femora and tibiae uniformly yellow without dark rings or with at most one inconspicuous brownish ring on midtibia. Abdomen. With setulose and microtrichose tergites, microtrichose pleural membrane without any setulae, and moderately narrow, microtrichose and setulose sternites without shiny spots. Female tergite 6 short, exposed beyond margin of tergite 5. Sternites 3–5 of female slightly longer than wide, sternites 4–6 with long anteromedial apodemes (Fig. 104). Male terminalia. Segments 6–8 sparsely microtrichose. Mesally-bent distal part of surstylus almost equal in length to basal portion; 3–4 prensisetae and numerous setulae; proctiger short; metaphallic plate long and pointed (Figs. 100, 103). Phallus long spinulose. Female terminalia. Oviscape microtrichose with bare dots at bases of setulae. Eversible membrane without visible scales. Aculeus long, basal portion almost twice as wide as cercal unit; the latter elongate oval, with long setulae. Three almost spherical spermathecae with short inverted necks (Fig. 105). Included species and distribution (Appendix 3, Fig. 134) Two species: Pseudotephritina cribellum (Loew, 1873: 134) (Stictocephala) (= Stictocephala cribrum Loew, 1873: 135) (Canada: Quebec; USA: Colorado; D.C.; Indiana; Iowa (new record); Kansas; Maryland; Michigan; Mississippi; Montana; Nebraska; New Mexico; New York; Ohio (new record); Pennsylvania; Texas; Virginia; Wyoming (new record)); P. inaequalis Malloch, 1932: 213 (Canada: Alberta (new record); USA: Arizona; California; Colorado; Indiana; Kansas; Nebraska; New Mexico; South Dakota; Texas; Utah (new record)) (Fig. 134). Biology Flies were collected mainly on poplar and willow, rarely on pear trunks; P. inaequalis was reared from cottonwood (Populus) bark in California (Steyskal, 1962). Determination Species can be determined using the following key adopted from Steyskal (1962). Key to species of Pseudotephritina 1. Propleural setulae black; second dorsocentral seta large. Male postabdomen with subequal prensisetae; metaphallic plate shorter than surstylus (Fig. 100) ……. P. cribellum (Loew). - Propleural setulae yellow; second dorsocentral seta small. Male postabdomen with basal prensiseta distinctly the largest; metaphallic plate as long as, or longer than, surstylus (Fig. 103) ...... P. inaequalis Malloch. Discussion Pseudotephritina is obviously a monophyletic genus. Its two species share 2 synapomorphies (the absence of an anterior orbital seta, and the presence of a long and pointed metaphallic plate), along with numerous symplesiomorphic characters. The most probable sister group of Pseudotephritina is Myennis. Species of these genera have non-speckled, microtrichose bodies with bare dots at the bases of setulae (less expressed in Myennis) and 2 shiny spots on the scutellum, apparent synapomorphies that are absent in both Pseudotephritis and Arborotites. In Pseudotephritina the wing venation remains unmodified (as in Pseudotephritis), whereas in both Myennis and Arborotites, either crossveins R-M and DM-Cu

548 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 97–105. Pseudotephritina. 97–100. P. cribellum. 97. Habitus, lateral view. 98. Head, dorsal view. 99. Mesonotum, dorsal view. 100. Epandrium, posterior view (from Steyskal, 1962). 101–105. P. inaequalis. 101. Wing. 102. Mesonotum and abdomen, dorsal view. 103. Epandrium, posterior view. 104. Female abdominal sternites 2–6. 105. Spermathecae.

549 BIOTAXONOMY OF TEPHRITOIDEA are more distal or the lobe of cell bcu is longer than in Pseudotephritis. The Arborotites + Myennis + Pseudotephritina lineage (the Myennis group) is supported by the modification of the intra-alar seta (in females, there is a cluster of shorter setulae instead of this seta).

Pseudodyscrasis Hernández, 1988 (Figs. 106-110, 135) Pseudodyscrasis Hernández-Ortiz, 1988: 183. Type species: scutellaris Wiedemann, 1830 (as “Pseudodyscrasis scutellaris (Wiedemann, 1830)”) (by original designation). Diagnosis Medium-sized (5.5–7.0 mm) flies, superficially similar to Myennis in wing pattern with oblique crossbands (however, arranged differently), uniformly colored legs, shiny scutellum, and gray microtrichose body without speckled pattern. The brief original description (Hernández-Ortiz, 1988: 183) needs to be refined, as one of the two originally included species has been removed to a new genus (see below). Redescription Head. As wide as or very slightly wider than thorax, 1.2 times as high as long and 1.1 times as wide as high. Eye 1.3 times as high as long. Frons 1.6 times as long as wide, in profile forming obtuse angle with dorsal portion of face, uniformly microtrichose; lateral margins parallel; anterior 0.8 of frons setulose, with bare median stripe reaching anterior margin. One lateroclinate ocellar seta. Two equally long orbital setae. Frontal setulae all black, 0.3 as long as orbital setae. Face slightly concave in profile, with conspicuous transverse furrow at ventral 0.16 and shallow antennal grooves. Clypeus receding, straight in profile, half as high as first flagellomere wide, completely microtrichose. Parafacial sparsely microtrichose, with bare spot at antennal base. ventral portion of facial ridge (to vibrissal angle) with 6–7 setulae. Anterior part of gena with row of 7–10 setulae reaching vibrissal angle. Gena 0.4–0.5 as high as eye. Genal seta well developed directly on genal ridge. Vertex without ridge; postocellar seta divergent, slightly longer than ocellar seta; vertical setae long, medial vertical seta twice as long as ocellar seta and 1.2 times than lateral vertical seta. Occiput sparsely microtrichose, without transverse dark mark or dots at bases of setulae. No paravertical setulae; 7–10 setulae in single postocular row; 3–4 occipital setulae laterally of ventral end of occipital suture and numerous setulae on high and convex postgena higher than level of occipital foramen. Posterodorsally curved supracervical setulae in two groups of 20–25 on each side separated by bare area. Hypostomal bridge short, with well developed gular seta and 3–5 shorter setulae at each ventrolateral corner. Antennae: distance between bases 1.2–1.7 times as wide as antennal socket. Scape rather large, but shorter than high, with 7–8 setulae dorsally. Pedicel wider than long, with blunt triangular medial lobe and shallow dorsal incision; lateral surface with one row of 18–20 marginal setulae, with 4 or 5 longest at ventral margin; 14–16 marginal setulae on medial surface forming 3 irregular rows; dorsal pedicellar seta slightly longer than pedicel. First flagellomere 1.5 times as long as wide, microtrichose. Two basal aristomeres. Terminal aristomere short pubescent. Proboscis stout, with large, rounded prementum that bears 14–16 setulae on each side. Labellum fleshy, large. Palpus very wide, widest at middle, with 20–25 rather short setulae on lateral and medial surfaces, and 8–10 setulae at ventral edge, some twice as long as lateral setulae, but shorter than palpus width.

550 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Thorax. Longer than wide, with gray and brown microtrichose areas, except shiny antepronotum, anterior margin of scutum, and scutellum. Postpronotal lobe with 1 seta and 10– 12 setulae. Proepisternum with group of 8–10 long posterodorsally directed setulae on dorsal portion and 1 weak proepisternal seta on ventral portion, shorter than these setulae. Prosternum slightly higher than wide, with 2 setulae at each side. Scutum with conspicuous dorsocentral row of setulae gradually becoming longer towards posterior margin; 1 row of acrostichal setulae, in posterior third somewhat scattered forming 2 rows; and 2–3 irregular rows of intra- alar setulae; 1 acrostichal, 3–4 dorsocentral, 1 intra-alar, 1 postalar and 1 supra-alar setae, all postsutural; other dorsocentral setulae less than half as long as longest seta. Bases of setulae with inconspicuous darker spots. Scutellum shiny. Subscutellum 0.2 times as long as scutellum, densely microtrichose. Notopleural triangle gray microtrichose on ventral portion, with velvety-black area on dorsal portion. Pleural sclerites thick gray microtrichose, anepistenum with brown round spots at bases of setulae. Wing. As in Figs. 106 and 107. Tegula unmodified, with 10–12 setulae and 2 marginal setae. Costal vein as described for Pseudotephritis; apical third of section between humeral break and apex of subcostal vein with 25–30 equally thickened setulae or thickened setulae alternated with 1–2 thin setulae on anterodorsal surface. Subcostal vein almost straight, gradually bent anteriorly in apical third. Vein R1 slightly bent anteriorly, bare before apex of vein Sc, apically with 10–12 setulae both dorsally and ventrally. Veins R2+3 and R4+5 slightly sinuous. Costal cell

1.2 times as long as pterostigma. Crossvein R-M perpendicular to vein R4+5, situated opposite middle of pterostigma, half as long as crossvein DM-Cu; cell br slightly widened at apex. Vein M: penultimate section twice as long as crossvein R-M and as long as crossvein DM-Cu. Crossvein BM-Cu somewhat oblique to vein M, not broken. Crossvein DM-Cu oblique to vein

M, not conspicuously sinuous. Vein Cu2 with posterodistal lobe of cell bcu twice as long as section between fork of vein Cu and elbow of vein Cu2. Posterodistal lobe of cell bcu triangular, slightly shorter than vein A1+Cu2. Basal cells (bc, c, br, bm and bcu) without dark spots or streaks isolated from veins by white borders. Calypteres as in Pseudotephritis, dorsal calypter with long white fringe. Halter not modified. Legs. Not thickened, sparsely microtrichose, as described for Myennis, except as follows. Midcoxa with row of 2 long setae and 3–4 shorter setulae on eucoxa and 10–12 setulae on disticoxa. Midtibia with one ventroapical seta twice as long as other thickened setae at apex and as long as tibial width; dorsal preapical seta not differentiated from setulae. Tarsi and their vestiture (including white brushes of fore and hindtarsomere 1) as in Pseudotephritis and Myennis. Abdomen. With tergites gray microtrichose and uniformly black setulose, pleural membrane microtrichose without any setulae and moderately narrow; sternites microtrichose and setulose. Tergite 1 partially brown microtrichose, tergite 4 and 5 with lateral spots of brown microtrichia. Female tergite 6 hidden underneath margin of tergite 5. Sternites 4–6 of female with anteromedial apodemes. Male terminalia. Segments 6-8 sparsely microtrichose. Epandrium with slightly curved, but not concave surstylus with mesally curved distal portion almost as long as basal portion of surstylus, with 3 long prensisetae and numerous setulae, proctiger short, metaphallic plate short, bluntly pointed at apex (Fig. 109). Phallus spinulose. Female terminalia. Oviscape sparsely microtrichose, long. Aculeus very long and narrow,

551 BIOTAXONOMY OF TEPHRITOIDEA

Figs. 106–117. Pseudodyscrasis, Neodyscrasis, n. gen. and Dyscrasis. 106–110. P. scutellaris. 106. Female habitus, lateral view (modified from Hendel, 1909 b). 107. Male wing. 108. Female mesonotum and abdomen, dorsal view. 109. Epandrium, posterior view (from Hernández, 1998). 110. Spermathecae. 111–114. N. steyskali. 111. Head, lateral view. 112. Wing. 113. Thorax and abdomen, dorsal view. 114. Epandrium, posterior view (from Hernández, 1998). 115–117. D. hendeli. 115. Epandrium, posterior view. 116. Female abdomen, ventral view. 117. Spermathecae (2 of 3).

552 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) more than half as long as preabdomen; cercal unit elongate oval, with long setulae. Three almost hemispherical spermathecae with long inverted necks (Fig. 110). Included species and distribution (Appendix 3, Fig. 135) Pseudodyscrasis scutellaris (Wiedemann, 1830) (Trypeta) (Mexico: D.F.; Hidalgo; México; Morelos; Veracruz). Hernández (1988) originally included in Pseudodyscrasis two species, P. scutellaris and P. steyskali Hernández (see below). Discussion Pseudodyscrasis and the two following genera form a group within the Myennidini, characterized by a low clypeus, large palpus, 2 equally long orbital setae, velvety-black posterior intra-alar area, more than two dorsocentral setae, a shiny scutellum, a microtrichose postscutellum with velvety-black lateral spots, non-banded legs, apically widened cell bm, more proximal position of crossvein R-M (at midlength of cell dm) and spermathecae with long necks. These numerous synapomorphies clearly show its monophyly. Pseudodyscrasis scutellaris fits the ground plan of the tribe more closely, at least in its completely microtrichose scutum, one supra-alar seta and basal wing cells without isolated dots and streaks, whereas the other two species of the group form a clearly monophyletic lineage. P. scutellaris possesses several autapomorphies which differentiate it from other myennidines: the setulose prosternum, ventrally setulose vein R1 and long aculeus. Including the two species originally assigned to the genus made Pseudodyscrasis paraphyletic and morphologically heterogeneous. For this reason, we remove P. steyskali Hernández to a separate genus (see below).

Dyscrasis Aldrich (Figs. 115-117, 118-119, 135) Dyscrasis Aldrich, 1932: 7. Type species: Dyscrasis hendeli Aldrich, 1932 (by original designation). Curran, 1934b: 279; Aczél, 1951: 429; Steyskal, 1961: 404; 1965: 644; 1987: 807. Description Aldrich (1932) briefly described the genus; that description must be completed as follows: Medium-sized flies (4.0–6.0 mm) superficially resembling Myennis in gray microtrichose body with shiny scutellum and spots on scutum and abdominal tergite 4, wings with yellow oblique crossbands and crossvein R-M situated proximaly to apex of vein R1, and non-banded legs. Dyscrasis differs from all other myennidines in having presutural acrostichal, dorsocentral and supra-alar setae. Head. As wide as thorax, 1.3 times as high as long and 1.8 times as wide as long. Eye almost as long as high. Frons as long as wide; lateral margins parallel; in profile forming right angle with dorsal portion of face; posterior half, including ocellar triangle and long vertical plates, microtrichose, with 4–6 setulae anteriorly, whereas anterior half shiny and bearing 18–20 setulae. One strong proclinate ocellar seta as long as frons. Two strong reclinate orbital setae; anterior at middle of frons, as long as posterior one. Frontal setulae 0.1–0.2 times as long as orbital setae. Face receding below antennae, protuberant at ventral margin. Clypeus small, less than half of first flagellomere width, completely hidden in peristomal cavity. Parafacial

553 BIOTAXONOMY OF TEPHRITOIDEA microtrichose with bare spot on dorsal half, concave. Facial ridge bare. Anterior part of gena with row of 4–6 dorsoclinate setulae reaching vibrissal edge. Gena 0.4 times as high as eye. Genal seta well developed, shifted posterior to genal ridge (border of gena and occiput). Vertical ridge slightly rounded, subacute; postocellar seta divergent, 0.3 times as long as ocellars; vertical setae very long, lateral vertical seta 0.5 times as long as medial vertical seta. Occiput white microtrichose, concave on dorsal portion, without dark marks. One paravertical setula or none; 8–15 setulae in single postocular row. Supracervical setulae not examined. Postgena moderately long setulose, with postgenal seta as long as genal seta, ventral to posterior tentorial pit. Hypostomal bridge short and bare. Antennae: distance between bases as wide as antennal socket. Scape short. Pedicel wider than long, with blunt triangular medial lobe and shallow dorsal incision; lateral surface with one row of 18–20 marginal setulae, with 4–5 longest at ventral margin; 15-20 setulae on medial surface forming 2–3 irregular rows; dorsal pedicellar seta almost as long as pedicel, shorter than longest ventral setulae. First flagellomere 1.5–1.6 times as long as wide, microtrichose. Two visible basal aristomeres. Terminal aristomere short pubescent. Proboscis stout, with large, rounded prementum bearing 12–15 setulae on each side. Labellum fleshy, large. Palpus very wide, widest at middle, with 10–12 setulae on lateral and medial surfaces much shorter than palpus width, and 8–10 setulae at ventral edge, mostly shorter than palpus width, and 4–5 setae as long as palpus width. Thorax. Gray microtrichose with brown microtrichose dots at bases of setae. Antepronotum, anterior margin of scutum, supra-alar areas anterior and posterior to transverse suture and entire dorsal surface of scutellum shiny. Postpronotal lobe with 1 seta and 5–7 setulae. Proepisternum with 4–8 long posterodorsally directed seta-like setulae on dorsal portion and 1 proepisternal setula-like seta on ventral portion. Prosternum subquadrate, without setulae. Scutum without setulae, with 1+4 (presutural and postsutural) acrostichals, 1+4 dorsocentrals, 1 intra-alar, 1 postalar and 1+2 supra-alar setae. No sexually dimorphic intra-alar setulae. Bases of setulae with spots of brown microtrichia. Scutellum convex, with 2 pairs of scutellar setae and without setulae, bare. Subscutellum 0.25 as long as scutellum. Notopleural triangle gray microtrichose on ventral portion and velvety-black on posterodorsal 0.33, with ochreous-gold medial portion. Anepistenum gray microtrichose, with brown microtrichose stripes on dorsal half and along ventral margin, without round spots; anepimeron gray and brown, katepisternum and meron completely brown, katatergite, anatergite and mediotergite gray microtrichose. Wing. As in Fig. 119. Tegula unmodified, with 4–5 setulae and 3–4 marginal setae. Costal vein as described for Pseudotephritis. Subcostal vein strongly sinuous: bent towards anterior margin at middle of costal sell, then back to vein R1 and again anteriorly. Vein Rs bare. Vein R1 slightly bent anteriorly, bare before Sc apex, apically with 18–20 dorsal setulae. Vein R2+3 slightly undulate. Vein R4+5 bare, ending before wing apex. Costal cell 1.1–1.2 times as long as pterostigma. Crossvein R-M oblique to vein R4+5, situated proximally of R1 apex, almost as long as crossvein DM-Cu; cell br conspicuously widened towards apex. Vein M: second section without stump vein; penultimate section 1.5–1.8 times as long as crossvein R-M and 1.4–1.5 times as long as crossvein DM-Cu; distal section straight, ending at wing apex. Crossvein BM- Cu perpendicular to vein M, broken at anterior end. Crossvein DM-Cu straight, almost perpendicular to vein M. Vein Cu2 Z-shaped, conspicuously receding in second section, with posterodistal lobe of cell bcu 4 times as long as section between fork of vein Cu and elbow of vein Cu2; vein A1+Cu2 reaching posterior wing margin. Posterodistal lobe of cell bcu triangular,

554 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Figs. 118–119. Dyscrasis hendeli. 118. Head, lateral view. 119. Male habitus, dorsal view.

twice as long as vein A1+Cu2, constricted at base. Cell bc, bases of cells c, br, bm and bcu with dark spots or streaks isolated by white border from veins (as in the tephritid Ceratitis MacLeay or the platystomatid Lenophila Guérin-Méneville). Vein A2 and alula as in Myennis. Dorsal calypter with long marginal fringe, 1.5–2 times as long as ventral calypter. Halter unmodified. Legs. Not thickened, sparsely microtrichose, as described for Myennis, except as follows: midcoxa with row of 7–8 long setae on eucoxa and 3–4 setae on disticoxa; midtibia with one ventroapical seta 3–5 times as long as other thickened setae at apex and twice as long as tibial width; dorsal preapical seta erect and thickened, but hardly 1.5 times as long as other setulae. Tarsi and their vestiture (including white brushes of fore and hindtarsomere 1) as in Pseudotephritis and Myennis.

555 BIOTAXONOMY OF TEPHRITOIDEA

Abdomen. Broad, with tergites at least partially setulose and microtrichose, pleural membrane broad without any setulae, and sternites moderately narrow, microtrichose and setulose. Tergite 4, and often tergites 2 and 3 with shiny spots laterally; male tergite 5 vestigial, shiny; female tergite 6 hidden underneath margin of tergite 5. Gray microtrichose area broken with dark brown spots at bases of setulae. Sternites 4–6 of female with anteromedial apodemes (Fig. 116). Male terminalia. Segments 6-8 almost shiny. Epandrium with slightly curved surstylus with mesal lobe slightly shorter than basal portion of surstylus, with 2 prensisetae and numerous setulae; proctiger short. Metaphallic plate moderately developed, parallel-sided, narrowed into nipple-shaped apex (Fig. 115). Phallus long spinulose. Female terminalia. Oviscape short, sparsely microtrichose. Aculeus comparatively short; cercal unit elongate oval, with moderately long setulae. Three almost spherical spermathecae with long inverted necks (Fig. 117). Included species and distribution (Appendix 3, Fig. 135) Dyscrasis hendeli Aldrich, 1932: 8 (USA: Arizona; Texas; Mexico: Compeche; D.F.; Durango; México; Morelos; Yucatán).

Neodyscrasis Kameneva and Korneyev, n. gen. (Figs. 111-114,135) Type species: Pseudodyscrasis steyskali Hernández, 1988. Gender: Feminine. Diagnosis Medium-sized flies (6.0–7.5 mm) similar to Dyscrasis in having shiny supra-alar spots on scutum, 2 supra-alar setae, apically broadened cell br, and similar dark brown spotted pattern of basal cells, differing in having numerous setulae, but no presutural setae on scutum and non- sinuous vein Sc. Description Head. As wide as thorax, 1.3 times as high as long and 1.6 times as wide as long. Eye 1.6 times as high as long. Frons slightly longer than wide; lateral margins parallel; in profile forming obtuse angle with dorsal portion of face; frons surface sparsely microtrichose, except ocellar triangle, vertical plates and narrow orbits densely white microtrichose, with numerous setulae; medial vitta bare, except anterior margin. One moderately developed ocellar seta. Two equal, moderately short, reclinate orbital setae; anterior orbital seta at posterior third of frons. Frontal setulae 0.25–0.3 times as long as orbital setae. Face slightly concave in profile, with unclear transverse furrow close to epistomal margin and inconspicuous antennal grooves. Clypeus as in Dyscrasis. Parafacial microtrichose with bare spot on dorsal 1/7. Facial ridge with 2–3 setulae above vibrissal corner. Anterior part of gena with row of 7–8 mostly proclinate setulae reaching vibrissal edge. Gena 0.3 times as high as eye. Genal seta well developed, posterior to genal ridge. Vertex moderately rounded; postocellar seta divergent, 1.1 times as long as ocellars; vertical setae moderately long, lateral vertical seta 0.8 times as long as medial vertical seta. Occiput white microtrichose, flattened or slightly convex, but not concave, with brown dots at bases of setulae. One paravertical setula or none; 6–8 setulae in single postocular row. Three to four occipital setulae lateral to suture and 4–5 setulae at level of occipital foramen. Two groups of 12-15 fine, white, dorsoclinate supracervical setulae on each side separated by bare area. Postgena moderately long and densely setulose to level of dorsal margin of foramen; postgenal

556 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae) seta ventral to posterior tentorial pit, as long as genal seta. Hypostomal bridge 4 times as wide as high, bare. Antennae: distance between bases as in Dyscrasis. Scape short, with dense black setulae exceeding length of scape. Pedicel as described for Dyscrasis, but more densely setulose; dorsal pedicellar seta not differentiated in examined specimens. First flagellomere 1.5–1.6 times as long as wide, microtrichose. Two basal aristomeres. Terminal aristomere bare. Proboscis as in Dyscrasis. Palpus wide, widest at middle, with 12–15 rather short setulae on lateral and medial surfaces, and 6–7 setulae at ventral edge, shorter than palpus width. Thorax. Vestiture as in Dyscrasis, except as follows. Postpronotal lobe with 1 seta and 10–12 setulae. Proepisternum with 8–10 posterodorsally directed setulae on dorsal portion; 1 setula- like proepisternal seta on ventral portion. Prosternum wider than high, without setulae. Scutum with conspicuous dorsocentral row of setulae gradually enlarged to setae posteriorly; 2 pairs of more or less regular rows of acrostichal setulae and 1–2 irregular intra-alar rows, also gradually becoming longer towards posterior margin; 0–2 presutural and 2–3 postsutural intra-alars, 0–1 presutural and 2 postsutural supra-alar setae and following setae only posterior to transverse suture: 5–8 acrostichals, 5–6 dorsocentrals and 1 postalar. Number of presutural setae varies among specimens (only those more than half as long as longest seta in row considered as setae). Bases of setulae with spots of brown microtrichia. Scutellum as in Dyscrasis. Subscutellum 0.25 times as long as scutellum, densely microtrichose. Notopleural triangle gray microtrichose on ventral portion and velvety black on dorsal portion. Pleura thick gray microtrichose, anepistenum with brown round spots at bases of setulae. Wing. As in Fig. 112. Tegula unmodified, with 3–4 setulae and 2–3 marginal setae. Costal vein as described for Pseudotephritis; apical third of section between humeral break and subcosta apex with 8–10 thickened setulae on anterodorsal surface; thickened setulae alternated with 1– 2 thin setae on anterodorsal surface, from apex of vein Sc to level of crossvein DM-Cu.

Subcostal vein almost straight, gradually bent anteriorly on apical third. Vein R1 slightly bent anteriorly, bare before Sc apex, apically with 18–20 dorsal setulae. Veins R2+3 and R4+5 as in

Dyscrasis. Costal cell 1.9 times as long as pterostigma. Crossvein R-M oblique to vein R4+5, situated at middle of pterostigma, half as long as crossvein DM-Cu; cell br widened towards apex. Vein M as in Dyscrasis; penultimate section 1.3 times as long as crossvein R-M and 0.7 times as long as crossvein DM-Cu. Crossvein BM-Cu almost perpendicular to vein M, not broken.

Crossvein DM-Cu sinuous, oblique to vein M. Vein Cu2 with posterodistal lobe of cell bcu twice as long as section between fork of vein Cu and elbow of vein Cu2. Posterodistal lobe of cell bcu triangular, as long as vein A1+Cu2. Cell bc, bases of cells c, br, bm and bcu with dark spots or streaks isolated from veins by white borders (as in Dyscrasis). Calypteres and halter as in Dyscrasis. Legs. As described for Dyscrasis. Abdomen. Wide, with tergites setulose and microtrichose, pleural membrane without setulae and sternites as in Dyscrasis. Tergites gray microtrichose, with dark brown spots at bases of setulae, but no shiny areas present. Female tergite 6 hidden underneath tergite 5 margin. Sternites 4–6 of female with apodemes. Male terminalia. Segments 6-8 subshiny. Epandrium as on Fig. 114, surstylus with apical portion (distal to prensisetae) longer than basal portion; 2 extremely long, claw-shaped prensisetae and additional setulae. Proctiger short. Metaphallic plate long, parallel-sided, narrow, gradually tapered and apically pointed into short apex (Fig. 114). Phallus long spinulose.

557 BIOTAXONOMY OF TEPHRITOIDEA

Female terminalia. Similar to those in Dyscrasis. Oviscape short, sparsely microtrichose. Included species and distribution (Appendix 3, Fig. 135) Neodyscrasis steyskali (Hernández-Ortiz, 1988: 8), comb. n. (Mexico: Chiapas, D.F., Durango, México, Morelos, Michoacán; Guatemala; Honduras; Venezuela) (see Kameneva, 2004, as Pseudodyscrasis steyskali, for distribution). Discussion The new genus is the sister-group of Dyscrasis. Both genera share numerous synapomorphic and symplesiomorphic characters (see diagnosis above). However, Dyscrasis possesses certain autapomorphies that strongly differentiate it from all Myennidini. Despite the monophyly of the Dyscrasis + Neodyscrasis lineage, N. steyskali and P. scutellaris share only a few symplesiomorphies, are very different in their appearance and diagnostic characters and do not form a monophyletic group. We therefore consider them to belong in different genera.

3. PHYLOGENETIC ANALYSIS OF THE MYENNIDINI The list of characters used in the analysis appears in Appendix 2, and the character matrix is in Table 1. Trees were calculated using both the HENNIG86 and PAUP 4.0 programs. One analysis with HENNIG86 in which no weight was assigned to all the characters was run. With the i.e.* (implicit enumeration) option HENNIG86 generated 156 cladograms (length 70, consistency index 57, retention index 81). Some examples of these trees are shown in Fig. 120. Two constant lineages appear on all the trees: the first includes all species of Callopistromyia, Oedopa, Stictoedopa, Paroedopa, and Stictomyia, while the second includes Pseudotephritis (except P. corticalis) and all species of Myennis, Arborotites, Pseudotephritina, Dyscrasis, Pseudodyscrasis and Neodyscrasis. The position of Pseudotephritis corticalis varies from tree to tree. It appears as the sister group either to the second of the above lineages (Fig. 120, trees 1, 2, 5, 7) or to the first (Fig. 120, trees 4, 6, 8), or in some cases in an unresolved trichotomy with both main lineages (Fig. 120, tree 3). The positions of many other terminal taxa also differ within the two main branches, but the following monophyletic groups are at least supported by all the analyses: Oedopa capito + O. ascriptiva¸ Stictomyia longicornis + S. punctata, Ulidiotites dakotana + Namibotites argentata¸ Pseudotephritis (except P. corticalis), Myennis, Pseudotephritina and Dyscrasis + Pseudodyscrasis + Neodyscrasis (henceforth “Dyscrasis group”). All the analyses show the following sister-group relationships: Pseudotephritina with the Dyscrasis group; Myennis with Pseudotephritina + Dyscrasis group; and Arborotites with Myennis + Pseudotephritina + Dyscrasis group. A Nelson consensus tree was obtained from 156 i.e.* trees (Fig. 121). When analyzed in HENNIG86, it has length 76, consistency index 0.52, retention index 0.77; the same tree, if rooted and analyzed in PAUP, has length 75, consistency index 0.5333 and retention index 0.7941 (the numbers are truncated to two decimals in HENNIG86 and to four in PAUP). These differences are due to unresolved basal polytomy including the outgroup in the HENNIG86 tree, which was corrected before importing it into PAUP. The monophyletic lineages found in the analysis with the implicit enumeration option, are also present in the PAUP tree. To resolve some of the polytomies and inconsistencies apparently due to incomplete morphological data and/or numerous homoplasies, successive weighting techniques were applied with use of both HENNIG86 and PAUP. When applying implicit enumeration combined with the successive weighting technique in

558 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Table 1 Matrix of character distribution in the Myennidini 0000000001 1111111112 2222222223 33 1234567890 1234567890 1234567890 12 Outgroup 000000000? 000???0000 0???000??0 ?? Arborotites stuckenbergi 0000000101 0000001000 1001001001 20 Pseudotephritis vau 0000000001 0001000000 0001001012 20 Pseudotephritis approximata 0000000001 0001100000 0001001011 20 Pseudotephritis millepunctata 0000000001 0000100000 0001001011 20 Pseudotephritis corticalis 0000000001 0000010000 0—1001000 21 Callopistromyia annulipes 0000000002 000-010100 1—1011000 21 Callopistromyia strigula 0000000002 000-010100 0—1011000 2? Oedopa capito 0210000002 10100-0100 1—1111000 11 Oedopa ascriptiva 0210000002 10100-0100 1—1111000 1? Oedopa elegans ?010000002 ??100-0100 ?—1????0? ?? Stictoedopa ruizi 1010000002 ?010000001 1—1????0? ?? Paroedopa punctigera 1110000002 10100-0000 1—1?11000 1? Stictomyia longicornis 1112000002 1010—0000 1—101110? 1? Stictomyia punctata 1111000002 1010-10000 1—1011100 12 Ulidiotites dakotana 0010000000 101—0001 0—001100? 11 Namibotites argentata 0000000000 001—0101 1—0001000 00 Myennis sibirica 0000000110 0101001000 0100001001 21 Myennis tricolor 0000000110 0200101000 0100001001 21 Myennis octopunctata 0000000110 0200001000 0100001001 21 Myennis mandschurica 0000000110 0200101000 0100001002 21 Pseudotephritina cribellum 0000001111 0001100010 0101001001 21 Pseudotephritina inaequalis 0000001110 0001100010 0100001001 21 Pseudodyscrasis scutellaris 0000101010 0000101000 0011001001 22 Neodyscrasis steyskali 0000111010 0000101000 1010001000 22 Dyscrasis hendeli 0000211010 0000101000 1010001000 22

HENNIG86, nine shortest trees (length 329, consistency index 0.79, retention index 0.91) were found in the second cycle, and further cycles did not change it. A Nelson consensus tree was obtained from these trees (Fig. 122). When analyzed in HENNIG86, it also has length 329, consistency index 0.79, retention index 0.91; the tree with the same topology, when rooted and analyzed in PAUP, has length 70, consistency index 0.5714 and retention index 0.8235. The principal differences from the “unweighted” consensus tree are that Paroedopa is the sister- taxon of Stictomyia, and Oedopa, excluding O. elegans, is the sister-taxon of Paroedopa + Stictomyia; Stictoedopa is the sister-group of Ulidiotites + Namibotites; and the polytomy in Pseudotephritis (except P. corticalis) is resolved, whereas in Myennis, there is unresolved polytomy among three terminal taxa. In the analysis with PAUP, characters 8 (the intra-alar area in female with cluster of setulae in addition to weak seta), 26 (the abdominal pleura setulose) and 27 (the metaphallic plate present) were assumed irreversible. A heuristic search (Hsearch) followed by contree and reweight index = ci commands after 3 repetitions produced an identical consensus tree (length 69, consistency index 0.5797, retention index 0.8294) (Fig. 123). In this tree, Namibotites has a more basal position on the Callopistromyia–Oedopa branch, manifesting its numerous differences from the Oedopa + Stictoedopa + Paroedopa + Stictomyia cluster, and also from

559 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 120. Possible phylogenetic relationships in the tribe Myennidini. Eight of 156 equally parsimonious trees resulting from HENNIG86 analysis with implicit enumeration (i.e.*) option.

560 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 121. Possible phylogenetic relationships in the tribe Myennidini. Nelson consensus tree of 156 equally parsimonious trees resulting from HENNIG86 analysis with i.e.* option and all the characters unweighted (“unweighted HENNIG86 analysis”). Character state changes represented by black bars are unique; those by gray bar are homoplasious; those by white bars are reversals. Ulidiotites, which looks superficially similar. The Dyscrasis group also has a more basal position, and Pseudotephritina appears as part of the Myennis cluster, with Myennis sibirica as its sister-group. In all three analyses, several more or less consistent clades were observed. The first major clade, which consists of Callopistromyia, Oedopa, Paroedopa, Stictoedopa, Stictomyia, Ulidiotites and Namibotites (the Callopistromyia –Oedopa clade), is supported by at least two synapomorphies (#10.2 and #26.1) in all the analyses, and additional synapomorphies in the two weighted analyses (#18.1 in the HENNIG86 analysis or #16.1, #32.1in the PAUP analysis). Of these synapomorphies, only the setulose abdominal pleura (#26.1) is unambiguous, although there is reversal in Namibotites (but see discussion below); the polarity of most of the other characters is not well substantiated. Another possible synapomorphy of this clade is the presence of three or more subequal, thickened apicoventral setae of the midtibia (#11.1); this character was not examined in

561 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 122. Possible phylogenetic relationships in the tribe Myennidini. Nelson consensus tree of 9 shortest trees resulting from HENNIG86 analysis with ie* option and successive weighting technique (“weighted HENNIG86 analysis”). Character state changes represented by black bars are unique; those by gray bar are homoplasious; those by white bars are reversals. Stictoedopa ruizi and Oedopa elegans, and the plesiomorphic condition occurs in Namibotites; consequently all the analyses show the apomorphic state dispersed among several taxa as a homoplastic character, but this may be due to incomplete morphological data. Both species of Callopistromyia are placed at the base of the Callopistromyia – Oedopa clade; they differ from other taxa of the clade in having the plesiomorphic condition of the setulosity of vein R1; indeed, both species have extensively developed speckled body and wing patterns, which are believed to constitue the ground plan of the tribe and are more or less reduced in the other taxa of this clade. No synapomorphies for both Callopistromyia species have been found, and the genus is possibly paraphyletic. The Oedopa clade (Oedopa, Paroedopa, Stictoedopa, Stictomyia, Ulidiotites and Namibotites) is recognized as monophyletic in all three analyses based on three synapomorphies: high gena (#3.1, except in Namibotites); bare vein R1 (#13.1); and finely setulose phallus (#31.1, bare (#31.0) in Namibotites). Within this clade, monophyly is supported for Stictomyia based on two synapomorphies (#4.1 (only if treated as ordered) and #28.1), and for Oedopa (except O. elegans) based on at least one (#2.2 and possibly #25.1).

562 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 123. Possible phylogenetic relationships in the tribe Myennidini. Tree resulting from PAUP analysis with heuristic search option combined with consensus and reweight from consistency index (“weighted PAUP analysis”). Characters 8, 26 and 27 are irreversible. Character state changes represented by black bars are unique; those by gray bar are homoplasious; those by white bars are reversals.

The Paroedopa + Stictomyia cluster is supported as monophyletic in both weighted cladograms by the presence of a round black spot between the antennae (#2.1). Stictoedopa is supported as the sister-group of Paroedopa + Stictomyia in the weighted cladogram that resulted from the PAUP analysis (Fig. 123) by head shape (short, but high and wide, #1.1). The distribution of these taxa also supports their possible relationship: they all occur in drier and more southern areas than the related genera Callopistromyia and Oedopa. However, morphological data on Stictoedopa, Paroedopa and Oedopa elegans remain incomplete, and further study may change the topology of this cluster. The relationships of Ulidiotites and Namibotites require special mention. The type species of both genera are superficially very similar in having uniformly silver-gray microtrichose bodies, yellow legs without dark rings, and hyaline wings with vein R1 bare and vein M anteriorly bent. Some of these characters (#10.0, 20.1, and #24.0) appear as synapomorphies on cladograms calculated with HENNIG86 (Figs. 121, 122). The placement of Namibotites within the Oedopa clade requires further assumptions of reversal in most of the important synapomorphies (#3.0,

563 BIOTAXONOMY OF TEPHRITOIDEA

#11.0, #26.0). Furthermore, Namibotites occurs in South Africa and may be most closely related to some Palearctic groups, either Myennis or Pseudotephritis, rather than to the Oedopa cluster, which is restricted to the New World. We therefore reject the hypothesis of close relationships between Namibotites and Ulidiotites (and with the Oedopa cluster at all) and consider its relationship to any major cluster within the Myennidini unproven. The second major clade, supported by all three analyses, consists of Pseudotephritis (except P. corticalis), Arborotites, Myennis, Pseudotephritina, and the Dyscrasis group. However, its monophyly is poorly supported by only two weak synapomorphies: the presence of more than 2 prensisetae (#30.1) is subject to reversal or homoplasy, and the absence of hyaline marks in the brown area bordering crossvein R-M (#16.0) is a character of uncertain polarity (not shown on the cladogram). Pseudotephritis (exclusive of P. corticalis) is supported as a monophyletic group in all three analyses by one synapomorphy (#29.1). The sister-group relationship of P. approximata with P. millepunctata is supported by the shape of the apical band (#15.1) on some unweighted cladograms from the HENNIG86 analysis; whereas P. vau appears as the sister-group of

P. approximata on the weighted cladograms because of the long medial spot in cell r1 (#14.1). The characters supporting each of these hypotheses vary in all three species and are of dubious value, thus we consider the relationships among P. millepunctata, P. approximata and P. vau unresolved (Fig. 121). Arborotites, Myennis, Pseudotephritina and the Dyscrasis group form a monophyletic cluster in the two HENNIG86 analyses (Figs 121, 122), supported by the approximated crossveins (#17.1); this character varies in some species of Myennidini, and the monophyly of this clade should be therefore considered dubious. In the analysis with PAUP, Arborotites has a more basal position, forming an unresolved polytomy with Pseudotephritis and the Myennis + Pseudotephritina + Dyscrasis cluster (Fig. 123). The Dyscrasis group is well supported as monophyletic in all three analyses by at least three synapomorphies (#5.1, #23.1, and #32.2). Of these synapomorphies, additional dorsocentral setae (#5.1) and the basal position of the second crossband on the wing (23.1) do not occur in other Myennidini. The presence of additional supra-alar setae (#6.1) supports the monophyly of the Neodyscrasis +Dyscrasis cluster. Pseudodyscrasis is its sister group in all the analyses. However, the position of the Dyscrasis group on the cladogram of the tribe varies considerably in different analyses. In both HENNIG86 analyses (Figs 121, 122), the Dyscrasis group forms a cluster with Pseudotephritina supported by the presence of a velvety-black spot at the base of the intra-alar seta (#7.1). In the weighted PAUP analysis with the “irreversible” option applied to certain characters, the Dyscrasis group appears as the sister group of Myennis +Pseudotephritina. The monophyly of this cluster is based on three weak synapomorphies: the absence of microtrichia on the scutellum (#9.1), absence or strong reduction of dark tibial rings (#10.0), and short distance between crossveins R-M and DM-Cu (#17.1). A shiny scutellum occurs in many lineages of Ulidiidae and higher Tephritoidea, thus independent reduction of microtrichia in both the Myennis +Pseudotephritina and Dyscrasis groups is not unlikely. Reduction of the dark rings on the tibiae also occurred in the Oedopa cluster (Ulidiotites), whereas the distance between crossveins considerably varies within the tribe. Another monophyletic group of genera that share a single apomorphy (#8.1) includes

564 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 124. Possible phylogenetic relationships in the tribe Myennidini. Tree manually modified from the “weighted PAUP analysis” tree with regards to objections to certain results of parsimony analysis. Characters 8, 26 and 27 are irreversible. Character state changes represented by black bars are unique; those by gray bar are homoplasious; those by white bars are reversals.

Myennis, Pseudotephritina and, possibly, Arborotites (henceforth, the “Myennis group”); the latter appears as a separate clade with character #8.1 as the result of homoplasy. The South African Arborotites retains characters that are symplesiomorphies with Pseudotephritis and Callopistromyia (speckled body coloration, microtrichose scutellum, tibiae with dark rings), whereas Myennis and Pseudotephritina share a partially shiny scutellum (#9.1), more distal position of the second crossband of the wing (#22.1), and a speckled body pattern reduced and replaced with round spots at bases of setulae or completely reduced on certain abdominal tergites (#24.0) (Fig. 124). All the analyses, however, show the Myennis group as non-monophyletic, with the Dyscrasis group arising within it. These computer-generated hypotheses are based on highly homoplastic characters (see above) and require the further assumption that character state #8.1 appears

565 BIOTAXONOMY OF TEPHRITOIDEA independently in Arborotites and Myennis + Pseudotephritina and disappears in the Dyscrasis group as reversal. This character, however, does not occur elsewhere in the Ulidiidae, and the latter hypothesis, in our opinion, is less probable than the parallel loss of dark coloration and scutellar microtrichia in the Myennis group and the Dyscrasis group. We consider them as separately evolved from a common, Pseudotephritis-like ancestor. In this case the basal branching of the Myennidini cladogram must be polytomic (Fig. 124). All the analyses support the monophyly of Pseudotephritina (#19.1) and of the cluster Myennis octopunctata + M. mandschurica + M. tricolor (#12.2), but the relationships of these taxa and M. sibirica are not well understood. M. octopunctata, M. mandschurica, M. tricolor and M. sibirica share a broadly darkened base of the costal cell (#12.1/12.2), which is a possible synapomorphy for Myennis, whereas Pseudotephritina and M. sibirica have a long medial spot beginning in cell r1 (#14.1). However, the polarity of the latter character is unclear. We consider the relationships among these three clades unresolved (Fig. 124). The position of Pseudotephritis corticalis within the Myennidini requires special mention, because in the unweighted trees from the HENNIG86 analysis it appears as the sister taxon either of the cluster of Pseudotephritis + Myennis group + Dyscrasis group or of the Oedopa cluster (Fig. 120). All of the consensus trees, both unweighted and weighted, leave it in an unresolved polytomy with these two clusters (Figs. 121–123). Indeed, it shares the speckled body pattern and single large midtibial seta (plesiomorphy or unclear polarity) with other Pseudotephritis, Arborotites, and Callopistromyia; it has a head shape and maculose wing pattern similar to Callopistromyia (polarity unclear), but its tibial pattern is like that in other Pseudotephritis (polarity unclear). If the maculose wing pattern is an apomorphy, then P. corticalis is the sister-group of the Oedopa cluster. If it is a plesiomorphic state, then P. corticalis must occupy a very basal position in the cladogram of the tribe. The distribution range (deciduous forest latitude zone and mountain areas south of it), which is unique in the Myennidini and rare in the Ulidiidae and higher Tephritoidea, may indirectly support such a basal position. Until new evidence is available, we consider this taxon in the basal unresolved polytomy with five other clusters (Fig. 124) The last cladogram (Fig. 124) was modified manually, mainly from the cladogram obtained with PAUP, with regard to certain objections to the results of parsimony analysis (see above) based upon the existing character set, which did not allow resolution of inconsistencies caused by homoplasies and incomplete morphological data. The main features of such an “intuitive” tree is that Namibotites is excluded from the Oedopa cluster and the Dyscrasis group is excluded from the Myennis lineage. Both are placed in unresolved basal polytomy. The tree (length 76; consistency index 0.5263; retention index 0.7895) is longer and less consistent than the consensus trees that resulted from the weighted PAUP and HENNIG86 analyses, but it is almost as short and consistent as the consensus tree from the unweighted HENNIG86 analysis, showing rather high probability of the “intuitively” hypothesized relationships.

ACKNOWLEDGMENTS We wish to express our gratitude to the following persons who kindly loaned or gifted us important material for this study: D. Azuma (ANSP); R.-E. Contreras-Lichtenberg (NHMW); J.M. Cummings (CNC); A. Freidberg (TAUI); D. Grimaldi (AMNH); A.L. Norrbom, N. Woodley, W.N. Mathis, L. Rodrigues and D.G. Furth (USNM); A.L. Ozerov and A.I. Shatalkin (ZMUM); L. Papp (HNHM); P.D. Perkins (MCZ); H. Wendt and W. Mey (ZMHB); C.D. Young (CMP); M. Zumbado (INBio).

566 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

We greatly appreciate the invaluable help from G.L. Bush, B. Merz, A. Freidberg, L. Papp, M. Zumbado, A. Herrera, F.C. Thompson and A.L. Norrbom, who made possible our visits to Tel Aviv, Budapest, Geneva, Washington, Santo Domingo, Vienna and Berlin in 1999–2002. A. Pont kindly supplied us with copies of Coquebert’s papers and his comments on some nomenclatural problems. B. Merz sent us a copy of Giglio-Tos’s papers and edited a translation of a French diagnosis. We are grateful to three anonymous reviewers and A. Freidberg for reviewing the manuscript and for their important comments. Special thanks to our son, Severin, and our parents for their patience and care. This study was carried out in partial fulfillment of the requirements for a Ph.D. degree by the first co-author (EPK) at the I.I. Schmalhausen Institute of Zoology, Kiev. In its final stage, this study was partially supported by travel grants from the Curtis Sabrosky Fund, Smithsonian Institution (Washington, D.C.) and the Ernst Mayr Fund, Cambridge University (Cambridge, Ma.) given to both co-authors, and by the Deutscher Akademischer Austauschdienst (DAAD, Bonn) stipend given to EPK, which provided a rare opportunity to study numerous specimens of more than 500 ulidiid species of almost all genera.

REFERENCES Aczél, M.L. 1951. Catálogo de la subfamilia americana “Pterocallinae” (Dipt. Acalypt. Otitidae). Acta Zoologica Lilloana 11: 397-433. Agassiz, L. 1846. Nomenclatoris zoologici index universalis. Jent et Gassmann, Soloduri. viii + 393 pp. Aldrich, J.M. 1905. A Catalogue of North American Diptera. Smithsonian Institution. Smithsonian Miscellaneous Collections. 46(2 [= No. 1444]). 680 pp. Aldrich, J.M. 1932. New Diptera, or two-winged flies, from America, Asia, and Java, with additional notes. Proceedings of the United States National Museum. 81 (9) [$2932], 28 pp. 1 pl. Anonymous. 1876. [Review of Rondani (1875) “Muscaria exotica Musaei civici januensis”] Bollettino della Società Entomologica Italiana 8: 177-184. Banks, N. 1914. A new ortalid . Proceedings of the Entomological Society of Washington 16: 138. Barraclough, D.A. 1995. An illustrated identification key to the acalyptrate fly families (Diptera: ) occurring in southern Africa. Annals of the Natal Museum 36 (October, 1995): 97-133. Barraclough, D.A. 1999. Amethysa Macquart, 1835, a genus of Ulidiidae (Diptera: Schizophora) errone- ously recorded from the Afrotropical Region. African Entomology 7(2): 285-286. Barraclough, D.A. 2000. A review of the Southern African Ulidiidae (Diptera: Schizophora), with descriptions of two new genera. Annals of the Natal Museum 41: 77-101. Becker, T. 1905. Cyclorrhapha Schizophora: Holometopa. pp. 1-273. In: T. Becker, M. Bezzi, K. Kertész, and P. Stein. Katalog der paläarktischen Dipteren. Band IV. Budapest. 328 p. Beschovski, V.L. 1972. A contribution to the dipteral fauna (Diptera, ) in the region of the Bulgarian Black-Sea Coast. Izvestiya na Zoologicheskiy Institut Bulgarska Akademiya na Naukite [= Bulletin de l’Institut Zoolgogique de l’Academie Bulgare] 34: 5-14. Bezzi, M. 1894. I ditteri del Trentino [conclusion]. Atti della Societá Veneto-Trentina di Scienza Naturali in Padova (2)1: 275-353. Bigot, J.M.F. 1852. Essai d’une classification générale et synoptique de l’ordre des insectes diptères. Annales de la Société Entomologique de France (2)10: 471-489. Bigot, J.M.F. 1885. [Les diagnoses de deux genres nouveaux de diptères appartenant à la famille des Ortalidae] Bulletin des Séances de la Société Entomologique de France (18): clxv-clxvi. Bigot, J.M.F. 1886. Diptères nouveaux ou peu connus. 29e partie. XXXVII. 1er. Essai d’une classification synoptique de groupe des Tanypezidi (mihi) et description de genres et d’espèces inédits. Annales de la Société Entomologique de France (6)6: 287–302. Bigot, J.M.F. 1892. Catalogue of the Diptera of the Oriental Region. Part III. Journal of the Asiatic Society of Bengal (n.s.) 61: 178–236.

567 BIOTAXONOMY OF TEPHRITOIDEA

Blanchard, É. 1846. Diptères. Athéricères. In: J.V. Audouin et al. [G.C.L.D. Cuvier] Le Règne . (q.v.). pl. 181. [Cited after Sabrosky, 1999] Brèthes, J. 1926. Desscription d’un noveau genre et nouvelle espèce d’Ulidiidae du Chile. Revista Chilena de Historia Natural 30: 187-188. Brues, C.T. Melander, A.L. and Carpenter, F.M. 1954. Classification of insects. (Revised ed.). Bulletin of the Museum of Comparative Zoology. Harvard University 108: 917 pp. Clements, D.K. 1998. Ulidiidae, pp. 122-123. In: P. Chandler (ed.). Checklists of Insects of the British Isles (New Series). Part 1: Diptera (Incorporating a List of Irish Diptera). London: Royal Entomo- logical Society. 234 pp. Clements, D.K. and Merz, B. 1997. The identity of ‘’ in Britain and its confusion with H. longistylata (Diptera, Ulidiidae). Dipterists Digest (4): 65-67. Clements, D.K. and Merz, B. 1998. Key to the genus Herina (Diptera, Ulidiidae) in Britain. Dipterists Digest (5): 55-67. Cole, F.R. 1969. The flies of Western North America. University of California Press. Berkeley and Los Angeles. 693 pp. Colless, D.H. and McAlpine, D.K. 1970. 34. Diptera (flies), pp. 656-740. In: The Division of Entomology. Commonwealth Scientific and Industrial Research Organisation, Canberra (spons.). The insects of Australia. A textbook for students and research workers. Melbourne University Press, Melbourne. 1029 pp. Colless, D.H. and McAlpine, D.K. 1991. Diptera (flies), pp. 717-786.In: The Division of Entomology. Commonwealth Scientific and Industrial Research Organisation, Canberra (spons.). The insects of Australia. A textbook for students and research workers. [Second edition.] Melbourne University Press, Melbourne 2: 543-1137. Coquillett, D.W. 1900. Notes and descriptions of Ortalidae. Journal of the New York Entomological Society 8: 21-25. Coquillett, D.W. 1910. The type-species of the North American genera of Diptera. Proceedings of the United States National Museum 37: 499-647. Coquillett, D.W. 1911. „Über die Nomenklatur der Acalyptratengattungen nach Th. Beckers Katalog der palaarktischen Dipteren, Bd. 4“. Wiener Entomologische Zeitung, 30(2/3): 62-64. Crampton, G.C. 1944. Suggestions for grouping the families of acalypterate cyclorrhaphous Diptera on the basis of the male terminalia. Proceedings of the Entomological Society of Washington 46: 152- 159. Cresson, E.T., Jr. 1906. Some North American Diptera from the South West. Transactions of the American Entomological Society 32: 279-288 + I pl. Curran, C.H. 1934a. The Diptera of Kartabo, Bartica District, British Guiana. Bulletin of the American Museum of Natural History 66 (art. 3): 287-533. Curran, C.H. 1934b. The families and genera of North American Diptera. New York, Ballou Press. 512 pp. Curran, C.H. 1938. New American Diptera. American Museum Novitates 975: 1-7. Desmarest, E. 1849. Ulidie, p. 753. In: d’Orbigny, C. Dictionnaire universel d’Histoire naturelle. Vol. 12. 816 pp. [Cited after Sabrosky, 1999] Desmarest, E. 1860. Table alphabétiques des noms vulgaires et scientifiques de tous les sujets décrits et figurés dans cette Encyclopédie. Annelés, 68 pp. In: J.Chenu. Encyclopédie d’histoire naturelle. Maresq, Paris. [Cited after Sabrosky, 1999] Enderlein, G. 1938. Die Dipterenfauna der Juan-Fernandez-Inseln und der Oster-Insel, pp. 643-680. In: K. Skottsberg (ed.). The Natural History of Juan Fernandez and Easter Island. Volume III. Zoology. 688 pp. Evenhuis, N.L. 1989. 63. Family Otitidae, pp. 479-481. Catalog of the Diptera of Australasian and Oceanian Regions. Honolulu and Leiden: Bishop Museum Press and E. J. Brill. 1155 pp. Fallén, C.F. 1810. Specim. entomolog. novam Diptera disponendi methodum exhibens. Lund, Berlingianis. 26 pp., 1 pl. Frey, R. 1921. Studien über den Bau des Mundes der niederen Diptera Schizophora nebst Bemerkungen über die Systematik dieser Dipterengruppe. Acta Societatis pro Fauna et Flora Fennica 48(3): 1- 246, 10 pls.

568 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Frey, R. 1964. Beitrag zur Kenntnis der ostasiatischen Plastystomiden (Diptera). Notulae Entomologicae 44: 1-19 + 3 pls. Ghesquière, J. 1947. [Ortaliscus], p. 691. In: P. Lepesme (ed.). Les insectes des palmiers. Paris, Chevalier. [iv] + 903 + [1] pp. Giglio-Tos, E. 1893. Diagnosi di nuovi generi e di nuove specie di Ditteri. IX. Bolletino del Musei di Zoologia ed Anatomia comparata della Universitá di Torino 8(158): 7-14. Giglio-Tos, E. 1895. Ditteri del Messico. Parte IV. Muscidae Calypteratae: Muscinae, Anthomyinae. Muscidae Acalypteratae: Scatophaginae, Helomyzinae, Tetanocerinae, Ortalinae, Ulidinae, Sapromyzinae, Trypetinae, Sepsinae, Tanypezinae, Psilinae, Chloropinae, Ephydrinae, Drosophilinae. Memorie della Reale Accademia delle Science di Torino. Serie Seconda 45: 74 p. Greve, L. 1998. 3.12. Family Otitidae, pp. 187-192. In: L. Papp and B. Darvas (ed.). Contributions to a Manual of Palaearctic Diptera. Volume 3. Budapest, Science Herald. 880 pp. Hardy, D.E. 1959. The Walker types of fruit flies Tephritidae (Diptera) in the British Museum collection. Bulletin of the British Museum (Natural History). Entomology 8: 159-242. Harriot, S.C. 1942a. New species of Otitidae from California (Diptera). The Pan-Pacific Entomologist. 18(1): 23-26. Harriot, S.C. 1942b. Notes on the genus Seioptera Kirby (Otitidae, Diptera). Journal of the New York Entomological Society. 50(June): 195–197. Harris, T.W. 1841. A report on insects of Massachusetts, injurious to vegetation. [1st edition]. Cambridge, Folsom, Wells and Thurston. viii + 459 pp. Heer, O. 1849. Die Insektenfauna der Tertiärgebilde von Oeningen und von Radoboj in Croatien. Zweite Theil: Heuschrecken, Florfliegen, Aderflügler, Schmetterlinge und Fliegen. Leipzig, W. Engel- mann. iv + 264 pp., 17 pls. Hendel, F. 1907. Nomina nova für mehrere Gattungen der Acalyptraten Musciden. Wiener Entomologische Zeitung 26: 98. Hendel, F. 1909a. Übersicht der bisher bekannten Arten der Pterocalliden (Dipt.). Deutsche Entomologische Zeitschrift. Beiheft. 1-84. Hendel, F. 1909b. Diptera, Fam. Muscaridae, Subfam. Pterocallinae In: P. Wytsman (ed.). Genera Insectorum. Bruxelles: Wytsman Verl. Fasc. 96. 50 pp., 4 pls. Hendel, F. 1909c. Beitrag zur Kenntnis der Ulidiinen (Dipt.). Wiener Entomologische Zeitung 28: 247- 270. Hendel, F. 1909d. Über die Gattung Euxesta Loew. (Dipt.). Annales [Historico-Naturales] Musei Nationalis Hungarici. 7: 151-172. Hendel, F. 1910a. Über acalyptrate Musciden. Wiener Entomologische Zeitung 29(2-3): 101-127. Hendel, F. 1910b. Diptera, Fam. Muscaridae, Subfam. Ulidiinae. In: Edit. P. Wytsman. Genera Insectorum. Bruxelles: Wytsman Verl. Fasc. 106 1-76, pls. 1-4. Hendel, F. 1910c. Über die Nomenklatur der Acalyptratengattungen nach Th. Beckers Katalog der paläarktischen Dipteren, Bd. 4. Wiener Entomologische Zeitung. 29: 307-313. Hendel, F. 1911. Über von Professor J. M. Aldrich erhaltene und einige andere amerikanische Dipteren. Wiener Entomologische Zeitung 30:19-46. Hendel, F. 1912. Neue Muscidae . Wiener Entomologische Zeitung 31(1): 1-20. Hendel, F. 1914a. Die Arten der Platystominen. Abhandlungen der (K.K.) Zoologisch-Botanischen Gesellschaft in Wien 8: 1-409. Hendel, F. 1914b. Neue amerikanische Dipteren. 2. Beitrag. Deutsche Entomologische Zeitschrift 1914(2): 151-176. Hendel, F. 1914c. Die Bohrfliegen Südamerikas. Übersicht und Katalog der bisher aus der neotropischen Region beschriebenen Tephritinen. Abhandlungen und Berichte des Koeniglichen Zoologischen und Anthropologisch-Ethnographischen Museums zu Dresden (1912) 14(3): 1-84. Hendel, F. 1916 Beiträge zur Systematik der Acalyptraten Musciden (Dipt.) Entomologische Mitteilungen 5: 294-299. Hendel, F. 1936a. Ergebnisse einer zoologischen Sammelreise nach Brasilien, insbesondere in das Amasonasgebiet, ausgefürt von Dr. H. Zerny. X. Teil. Diptera: Muscidae acalyptratae (excl. Chlo- ropidae). Annalen des Naturhistorischen Museums in Wien 47: 61-106.

569 BIOTAXONOMY OF TEPHRITOIDEA

Hendel, F. 1936b. 26. Ordnung der Pterygogenea (Dreissigste Ordnung der Insecta): Diptera oder Fliegen [part], pp. 1885-1980. In: W. Kükenthal and T. Krumbach (eds.). Handbuch der Zoologie 4, 2. Halfte, Insecta 3. Walter de Gruyter & Co. Berlin and Leipzig (1936-1937): 1729-1998. Hennig, W. 1939. 46/47. Otitidae (46. Otitidae und 47. Pterocallidae). In: E. Lindner (ed.). Die Fliegen der palaearktischen Region, 5(126-128). Stuttgart, E. Schweizerbart. 79 pp. Hennig, W. 1940. 45. Ulidiidae. In Lindner E. (ed.). Die Fliegen der paläarktischen Region, 5(133). Stuttgart, E. Schweizerbart. 34 pp. Hennig, W. 1952. Die Larvenformen der Dipteren. Eine Übersicht über die bisher bekannten Jugendstadien der zweiflügen Insekten. 3. Teil. Berlin, Akademie-Verlag. 628 + vii pp., 21 pls. Hennig, W. 1958. Die Familien der Diptera Schizophora und ihre phylogenetischen Verwandschafts- beziehungen. Beiträge zur Entomologie 8: 505-688. Hennig, W. 1973. Ordnung Diptera (Zweiflügler). Handbuch der Zoologie. Walter De Gruyter. Berlin. 4, 2. 2/31 (Lieferung 20): 1-337. Hering, E.M. 1956. Eine neue Myennis-Art (Dipt. Otitidae). Deutsche Entomologische Zeitschrift N. F. 3: 87-90. Hernández-Ortiz, V. 1988. Reconsideracion taxonomica del genero Dyscrasis Aldrich y la descripcion de Pseudodyscrasis gen. nov. (Diptera: Otitidae). Folia Entomológica Mexicana 74: 181-188. Hunter, W.D., Pratt, F.C. and Mitchell, J.D. 1913. The principal cactus insects of the United States. Bulletin of the United States Department of Agriculture Bureau of Entomology [n. ser.] 113: 1-71, 7 pls. International Commission on Zoological Nomenclature. 1985. International Code of Zoological Nomen- clature. Third Edition adopted by the XX General Assembly of the International Union of Biological Sciences. International Trust for Zoological Nomenclature, London. xx + 338 pp. International Commission on Zoological Nomenclature. 1999. International Code of Zoological Nomen- clature. Fourth Edition adopted by the International Union of Biological Sciences. International Trust for Zoological Nomenclature, London. xxix + 306 pp. Johnson, C.W. 1902. Remarks on Tephronota ruficeps and description of a new species. Entomological News 13: 143-144. Johnson, C.W. 1915. A new species of Pseudotephritis. Psyche 22: 49. Johnson, C.W. 1921. New species of Diptera. Memoirs of the Boston Society of Natural History 5: 11-17. Kameneva, E.P. 1996. A new species of the genus Timia (Diptera: Ulidiidae) from Tien-Shan mountains. Journal of the Ukrainian Entomological Society 2(3-4): 51-55. Kameneva, E.P. 1997a. Revision of the palaearctic species of Melieria R.-D. (Diptera, Ulidiidae, Otitinae). 1. The groups of species assigned to Hypochra Loew. Vestnik Zoologii (1996). 30(6): 19-46. Kameneva, E.P. 1997b. Status of Systata Loew (Diptera: Ulidiidae: Otitinae). Journal of the Ukrainian Entomological Society 3(1): 49-54. Kameneva, E.P. 2001. 76-77. Fam. Ulidiidae (Otitidae, Pterocallidae, Ortalidae) — picture-winged flies, pp. 151-165. In: P.A. Lehr (ed.). Key to Insects of the Far East Russia. Vol. VI. Diptera and Fleas. Part 2. Vladivostok, Dal’nauka. 639 + [2] pp. [In Russian] Kameneva, E.P. 2004. New records of picture-winged flies (Diptera: Ulidiidae) of Central America, including the description of new taxa. Studia dipterologica (2003) 10(2): 609-652. Kameneva, E.P. and Korneyev, V.A. 1994. Holarctic genus Pseudoseioptera Stackelberg (Diptera: Ulidiidae (= Otitidae)). Part I. Phylogenetic relationships and taxonomic position. Journal of the Ukrainian Entomological Society (1993) 1(2): 65-72. Korneyev, V.A. 1999. Phylogenetic relationships among the families of the superfamily Tephritoidea, pp. 3-22. In: M. Aluja, A.L. Norrbom (eds.). Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. Boca Raton – London – New York – Washington, CRC Press. 944 pp. Krivosheina, N.P. 1998. The redescription of the type specimens of Myennis mandschurica Hering, 1956 (Diptera, Otitidae). International Journal of Dipterological Research 9(2): 95–98. Krivosheina, N.P. and Kovalev, V.G. 1972. New data on flies of the genera Seioptera Kirby and Pseudoseioptera Stack. (Diptera, Otitidae). Izvestiya SO AN SSSR 5(1): 81–86 [in Russian].

570 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Krivosheina, N.P. and Krivosheina, M.G. 1995a. Euxesta stackelbergi sp. n. — new species of Ulidiidae (Diptera), discovered under the bark of Ulmus foliacea Gilig. in Turkmenistan. International Journal of Dipterological Research 6: 49-53. Krivosheina, N.P. and Krivosheina, M.G. 1995b. A revision of the genus Homalocephala (Diptera, Ulidiidae) of Russia. Russian Entomoogical Journal 4(1-4): 109-113. Krivosheina, N.P. and Krivosheina, M.G. 1997a. New data on representatives of the genus Physiphora (Diptera, Ulidiidae). Zoological Journal 76(4): 460-465. Krivosheina, N.P. and Krivosheina, M.G. 1997b. Revision of palaearctic species of the genus Myennis (Diptera, Otitidae) Zoological Journal 76(5): 628-632. Krivosheina, N.P. and Krivosheina, M.G. 1997c. Revision of palaearctic species of the genus Pseudotephritis Johnson (Diptera, Otitidae). Entomologicheskoe Obozrenie 76(3): 671-679. Krivosheina, M.G. and Krivosheina, N.P. 1997d. New data on palaearctic species of the genus Euxesta (Diptera, Ulidiidae), with a description of a new species Euxesta freyi. Zoological Journal 76(10): 1179-1184. Lioy, P. 1864. I ditteri distribuiti secondo un nuovo metodo di classificazione naturale [part]. Atti del Reale Istituto Veneto di Scienze, Lettere ed Arti (3) 9: 499-1352. Loew, H. 1862. Monographs of the Diptera of North America. Part I. Smithsonian Miscellaneous Collections. 6(1 [= No. 141]). xxiv + 221 pp. Loew, H. 1866. Empylocera, eine neue Ortaliden-Gattung. Berliner Entomologische Zeitschrift 10: 238- 240. Loew, H. 1868a. Die amerikanischen Ulidiina. Berliner Entomologische Zeitschrift (1867). 11: 283-326. Loew, H. 1868b. Die europaischen Ortaliden. Zeitschrift für die Gesammten Naturwissenschaften Halle 32: 1-11. Loew, H. 1873. Monographs of the Diptera of North America. Part III. Smithsonian Miscellaneous Collections. 11[= No. 256]. vii + 351 + XIII pp., 4 pls. Macquart, J. 1835 Histoire naturelle des insectes. Diptères. Tome deuxième. Paris, Roret. 703 or 710 pp., 12 pls. Malloch, J.R. 1931. Flies of the genus Pseudotephritis Johnson (Diptera: Ortalidae). Proceedings of the United States National Museum 79(34): 1-6. Malloch, J.R. 1932. New species and other records of Otitidae (Ortalidae), Piophilidae, Clusiidae, Chloropidae, and Drosophilidae from the Marquesas. In: Marquesan Insects. I.B.P. Bishop Museum Bulletin. Pacific Entomological Survey Publication I. Arts 1, 2, 11, 14: 205-223. Malloch, J.R. 1933. Fascicle 4. Acalyptrata. In: Diptera of Patagonia and South Chile. [London, British Museum]. 6: 177-386. Malloch, J.R. 1940. Otitidae and Phytalmiidae of the Solomon Islands (Diptera). Annals and Magazine of Natural History Ser. II. 6(July, 1940): 66-98 + 1 plate. Mamaev, B.M., Krivosheina, N.P. and Pototskaya, V.A. 1977. Keys to larvae of insects living in wood. Moscow, Nauka. 368 pp. [in Russian] McAlpine, J.F. 1977. A revised classification of the Piophilidae, including “Neottiophilidae” and “Thyreophoridae” (Diptera: Schizophora). Memoirs of the Entomological Society of Canada 103: IV + 66 pp. McAlpine, J.F. 1989. Phylogeny and classification of the , pp. 1397-1518. In: J.F. McAlpine (ed.) and D.M. Wood (coordinators). Manual of Nearctic Diptera. Vol. 3. Ottawa: Research Branch Agriculture Canada. 1989. Monograph No 32. i–vi + pp 1333-1581. Merz, B. 1996a. Systematik und Faunistik der Gattung Herina (Diptera, Otitidae) der Schweiz. Mitteilungen der Schweizerischen Entomologische Gesellschaft 69: 329-344. Merz, B. 1996b. Zur faunistik der Pyrgotidae, Platystomatidae und Ulidiidae (= Otitidae) (Diptera, Tephritoidea) der Schweiz mit spezieller Berücksichtigung von Otites Latreille. Mitteilungen der Schweizerischen Entomologische Gesellschaft 69: 405-416. Merz, B. 1998. 61. Ulidiidae (= Otitidae), pp. 242-243. In: B. Merz, G. Bächli, J.-P. Haenni and Y. Gonseth (eds). Diptera – Checklist. Fauna Helvetica. 1. Neuchâtel: CSCF und SEG: 369 pp. Osten Sacken, C.R. 1881a. [Diagnoses de cinq nouveaux genres de diptères exotiques de la division des Orthalidae.] Bulletin des Séances de la Société Entomologique de France 1881 (15): xcix-c.

571 BIOTAXONOMY OF TEPHRITOIDEA

Osten Sacken, C.R. 1881b. [Diagnoses de cinq nouveaux genres de diptères exotiques de la division des Orthalidae.] Bulletin de la Société Entomologique de France (6) 1: xcix-c. Page, R.D.M. 2001a. NEXUS Data Editor 0.5.0. Program. http://taxonomy.zoology.gla.ac.uk/rod/ rod.html. Page, R.D.M. 2001b. TreeView (Win32) 1.6.5. Program. http://taxonomy.zoology.gla.ac.uk/rod/rod.html. Ramos, T.C. 1997. TreeGardener 2.2. Program and manual. [Museu de Zoologia, Universidade de São Paulo, Brazil] Richter, V.A. 1970a. 60. Fam. Otitidae, pp. 123-130. In: G.Ya. Bey-Bienko (ed.). Key to species of European Territory of the USSR in five volumes. V. Diptera, Fleas. Second part. Leningrad, Nauka. 943 pp. [In Russian] Richter, V.A. 1970b. 61. Fam. Ulidiidae, pp. 130-132. In: G.Ya. Bey-Bienko (ed.). Key to species of European Territory of the USSR in five volumes. V. Diptera, Fleas. Second part. Leningrad, Nauka. 943 pp. [In Russian] Rohdendorf, B.B. 1964. Historical development of dipterous insects. Trudy Paleontologicheskogo Instituta. 100. Moscow, Nauka. 311 pp. [In Russian] Robineau-Desvoidy, J.B. 1826. [Names of family group and genera.] In: Académie Royale des Sciences de l’Institut de France, H.M.D. de Blainville (Rapporteur), Rapport sur les Myodaires du Docteur Robineau Desvoidy. Académie Royale des Sciences, Paris. 24 pp. Robineau-Desvoidy, J.B. 1830. Essai sur les myodaires. Mémoires Présentés par divers Savants a l’Académie Royale des Sciences de l’Institut de France 2(2). 813 pp. Rondani, C. 1856. Dipterologiae Italicae prodromus. Vol: I. Genera Italica ordinis dipterorum ordinatim disposita et distincta et in familias et stirpes aggregata. A. Stoschi, Parmae. 226 + [2] pp. Rondani, C. 1869. Ortalidinae Italicae collectae, distinctae et in ordinem dispositae [part]. Bollettino della Società Entomologica Italiana 1: 5-37. Rye, E.C. 1873. Insecta. Diptera. Zoological Record (1871) 8: 384-397. Sabrosky, C.W. 1946. Family names in the order Diptera. Proceedings of the Entomological Society of Washington 48: 163-171. Sabrosky, C.W. 1999. Family-group names in Diptera. Myia 10: 1-360. Schiner, I.R. 1864. Catalogus systematicus dipterorum Europae. Vindobonae [=Vienna], Societatis zoologico-botanicae. xii + 115 pp. Sharp, D. 1899. Insects. Part II. Hymenoptera continued (Tubulifera and Aculeata), Coleoptera, Strepsiptera, Lepidoptera, Diptera, Aphaniptera, Thysanoptera, Hemiptera, Anoplura. In: Harmer, S. F. and Shipley, A.E. (eds.) The Cambridge natural history. London, Macmillan and Co. Vol. 6. xii + 626 + 2 pp. Snow, W.A. 1896. [Ortalidae] Table of Genera, pp. 115-119. In: Williston, S.W. Manual of the families and genera of North American Diptera. (ed. 2). New Haven, J.T. Hathway. liv + 167 pp. Soós, Á. 1957. Neue Angaben die paläarktischen Otitiden (Diptera). Annales historico-naturales musei nationalis Hungarici (Series nova) 8: 389-399. Soós, Á. 1984. Family Otitidae (Ortalidae), pp. 45–59. In Soós, A., and Papp, L. (eds.). Catalogue of Palaearctic Diptera, Vol. 9, Micropezidae–Agromyzidae. Akadémiai Kiádo, Budapest and Elsevier Science Publishers, Amsterdam. 460 pp. Steyskal, G.C. 1952. Ulidiinae (Diptera: Otitidae) of Australasian Region. Occasional Papers of B.P. Bishop Museum Honolulu, Hawaii 20(15): 277-287. Steyskal, G.C. 1961. The genera of Platystomatidae and Otitidae known to occur in America North of Mexico (Diptera, Acalyptratae). Annals of the Entomological Society of America 54(3): 401-410. Steyskal, G.C. 1962. The American species of the genera Melieria and Pseudotephritis (Diptera: Otitidae). Papers of the Academy of Science, Arts, and Letters. Ann Arbor, Michigan. (1961) 47: 247-262. Steyskal, G.C. 1965. Family Otitidae, pp. 642-654. In: A. Stone, C.W. Sabrosky, W.W. Wirth, R.H. Foote and J.R. Coulson (eds.). A Catalog of the Diptera of America north of Mexico. Washington: United States Department of Agriculture. Handbook N 276. iv + 1696 pp. Steyskal, G.C. 1968. 54. Family Otitidae (Ortalidae; including Pterocallidae, Ulidiidae). In: P.E. Vanzolini, N. Papavero (eds.). A catalogue of the Diptera of the Americas south of the United States. São Paulo: Dept. Zool. Secretaria Agric. Fasc. 54. 31 pp.

572 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Steyskal, G.C. 1974. The genus Oedopa Loew (Diptera, Otitidae, Ulidiinae). Proceedings of the Entomo- logical Society of Washington. 76: 163-164. Steyskal, G.C. 1975. Genus Callopistromyia Hendel (Diptera: Otitidae). Entomological News (1974). 85(5-6): 147-148. Steyskal, G.C. 1977. Family Otitidae, pp. 165-167. In: M.D. Delfinado and D.E. Hardy (eds.). A catalog of the Diptera of the Oriental Region. Vol. 3. Suborder Cyclorrhapha (excluding Division ). Honolulu: The University Press of Hawaii. x + 854 pp. Steyskal, G.C. 1979. Biological, anatomical, and distributional notes on the genus Callopistromyia. Proceedings of the Entomological Society of Washington 81(3): 450-455. Steyskal, G.C. 1980. 44. Family Otitidae, pp. 575-577. In: R.W. Crosskey (ed.). A catalogue of the Diptera of the Afrotropical Region. London: British Museum (Natural History). 1437 pp. Steyskal, G.C. 1982. A key to the genera of the subfamily Otitinae of the Americas South of the United States (Diptera: Otitidae). Memoirs of the Entomological Society of Washington 10: 139-144. Steyskal, G.C. 1987. Otitidae. 63, pp. 799-808. In: J.F. McAlpine (ed.), B.V. Peterson, G.E. Shewell, H.J. Teskey, J.R. Vockeroth, D.M. Wood (cordinators). Manual of Nearctic Diptera. Vol. 2. Ottawa: Research Branch Agriculture Canada, 1987. Monograph No 28. i–vi + pp. 675-1332. Sturtevant, A.H. 1926. The seminal receptacles and accessory glands of the Diptera, with special reference to the Acalypterae. Journal of the New York Entomological Society (1925) 33: 195-215. Swainson, W. 1840. Diptera, pp. 366-380. In: W. Swainson and W.E. Shuckard. On the history and natural arrangement insects. London, Longman, Orme, Brown, Green & Longmans’s [&] J. Taylor, London. [x] + 406 pp. Swofford, D.L. 1998. PAUP*. Phylogenetic Analysis Using Parsimony (*and Other Methods). Version 4. Sinauer Associates, Sunderland, Massachusetts. Thompson, F.C. and Pont, A.C. 1994. Systematic database of Musca names (Diptera). A catalog of names associated with the genus-group name Musca Linnaeus, with information on their classification, distribution, and documentation. Theses Zoologiae. 20. Koenigstein, Koeltz Scientific Books. (1993). 221 pp. Vos de Wilde, B. 1935. Contribution a l’étude des larves de Diptères Cyclorraphes, plus spècialement des larves d’Anthomyies. Proefschrift. Amsterdam. 125 pp. [Cited after Ferrar, 1987] Westwood, J.O. 1840. Order XIII. Diptera Aristotle. (Antliata Fabricius. Halteriptera Clairv.), pp. 125- 154. In: J.O. Westwood. An introduction to the modern classification of insects. Synopsis of the genera of British Insects. Vol. 2. London, Longman, Orme, Brown, Green and Longmans. 587 pp. White, I.M., Headrick, D.M., Norrbom, A.L., and Carroll, L.E. 1999. Glossary, pp. 881-924. In: M. Aluja and A.L. Norrbom (eds.). Fruit Flies (Tephritidae): Phylogeny and Evolution of Behavior. Boca Raton—London—New York—Washington, CRC Press. 944 pp. Williston, S.W. 1908. Manual of North American Diptera. Ed. 3. New Haven, Conn. 405 p. Wulp, F.M. van der. 1898. Fam. Muscidae [part], pp. 385-416. In: F. Godman and O. Salvin (eds.). Biologia Centrali-Americana. Zoologia. Class Insecta. Order Diptera. 3 vols. Vol. 2. London, Taylor and Francis. (1899-1901). 489 pp., XIII pls. Zaitzev, V.F. 1982. Flies of the family Ulidiidae (Diptera) in the fauna of Mongolia. Nasekomye Mongolii. 8. Leningrad, Nauka: 422-453. Zaitzev, V.F. 1984. Family Ulidiidae. pp. 59-66. In: A. Soós and L. Papp (eds.). Catalogue of Palaearctic Diptera, Vol. 9, Micropezidae—Agromyzidae. Budapest and Amsterdam, Akadémiai Kiádo and Elsevier Science Publishers. 460 pp. Zuska, J. 1967. The Czechoslovak species of Ulidiidae (Diptera, Acalyptrata). Acta faunistica entomologica Musei Nationalis Pragae 12(130): 201-206.

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APPENDIX 1 Myennidini material examined for this study (taxa arranged alphabetically)1 Arborotites stuckenbergi: South African Republic: “Sarnia (Natal)”, 15.i.1912, & (Janse) (with label: “Stuckenbergia Steyskal / ectopa Steyskal”) (HNHM). Callopistromyia annulipes: Non-type material: Canada: Alberta: Champion, dead poplar bark, 14.ix.1961, 3 (, 8 & (FIS) (CNC); British Columbia: Robson, 5.vi.1952, 2.ix.1952, 12.ix.1952, 3 & (Foxlee) (CNC); USA: Alabama: Florence, Pollo study trap, &, 25.vii.1946 (USNM); Arizona: White Mts., 25.vii.1944, & (Parker and Lot) (USNM); Connecticut: Hamden, 14.iv.1939, & (Farland) (USNM); Georgia: Glassy Mt. / Rabun Co. 12.v.1953, & (Seago) (USNM); Idaho: Goodling, 25.vii.1944, 2 (, 5 &; Payette, 27.vii.1944, (, & (Peach) (USNM); Illinois: Champain, 1.vi.1953, & (J.F.McAlpine) (CNC); Chicago, & (USNM); Iowa: Ames, State Nursery, Malaise trap, 20.vi.1972, (, & (Pinger) (USNM); Indiana: Lafayette, on tree trunk, 5.vi–25.ix.1915, (, 3 & (Aldrich) (USNM); Louisiana: Melville, 11.v.1929, & (Schott) (USNM); Maine: Monmouth, 24.vi.1957, & (USNM); Bar Harbor, 16.vii.1958, & (Vockeroth) (CNC); Maryland: Talbot County, McDaniel (Wades Point), Malaise trap in salt marsh with flowering Baccharis, 19-21.ix.1986, 1 & (Steiner) (USNM); Massachusetts: Beverly, 18.9.1870, (, 3 & (USNM); Michigan: Traverce City, 16.vi.1943, 3 &; Bear Lake, 25.vi.1943, 2 &; Nottawa, 27.viii.1932, 29.viii.1934, 2 (; East Lansing, 19.viii.1942, 2 (, & (Sabrosky) (USNM); Mississippi: Lafayette Co., v–vi.1960, 2 (, 5 & (Hull) (CNC); Missouri: Columbia, Malaise trap, 4 pm-7 am, 7.vi.1967, & (Parker) (USNM); Nebraska: Lincoln, reared from Ailantus bark, 12.vii.1962, (, 3 & (Hanson) (USNM); North Carolina: “N.C.”, &; Tryon 22.vii.1941, 2 (, & (Melander) (USNM); Ohio: “Ohio”’ &; Athens, 1.ix.1931, & (USNM); Oxford, 05.1945, 2 (, 3 & (USNM); Tennessee: Great Smoky Mts. National Park, Greenbrier Cove, 2000', 18.v.1957, 3 & (Vockeroth); Utah: Salt Lake City, 1928, 15 (, 8 & (Cowie); Ogden, 3.iv.1943, ( (Knowlton and S. L. Wood) (USNM); Washington: Walla Walla, 2.v.1940, & (Lanchester); Yakima Co., 11.vii.1944, & (USNM); West Virginia: Hardy County, Lost River State Park, 3.ix.1978, 8 (, 18 & (Steyskal) (USNM); Wisconsin: Madison, 2.vii.1951, & (Snyder) (USNM); about 60 other specimens from various localities in New Jersey, New York, Pennsylvania, Virginia, D.C. and Maryland. Callopistromyia strigula: Type material: Syntype (: [USA]: “Georgia. Poppig” [green label], “2800”, “Pterocalla strigula Loew 1873 ( Dr. Enderlein det.”, “Typus” [red label of “Enderlein’s” type”], “Zool. Mus. Berlin” (ZMHB). Non-type material: Canada: Saskatchewan: Regina, 2.vii.1905, & (Willing) (USNM); USA: Illinois: Urbana / Ill., 8.viii.1902, (, & (Webster) (USNM); Iowa: Ames, State Nursery, Malaise trap, 25.vi.1972, (, 3 & (Pinger) (USNM); Kansas: Lawrene, 900 ft., May, & (Tucker); idem, 16.v.1897, 3 (, 3 & (USNM); Maryland: Mtg. Co., C and O Canal near Plummers Island, on fallen branch of Platanus, 31.viii.1989, ( (Norrbom and Steiner) (USNM); Michigan: Lapeer Co., Deerfield twp. 5.ix.1943, (, &, 11.viii.1946, (; Wayne Co., Grosse Ile. 31.v.1948, (, 26.ix.1958, & (Steyskal) (USNM); Minnesota: Rush City, 7.ix.1952, & (Sabrosky) (USNM); Nebraska: Lincoln, April (, & (USNM); Pennsylvania: Pinsleny, 24.vii.1907, & (USNM); South Dakota: Brookings, ( (Aldrich) (USNM); Minnesota: Rush City, 7.ix.1952, & (Sabrosky) (USNM);

1Labels of primary types are cited verbatim, with slash separating lines and back-slash separating label downside inscriptions.

574 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Utah: Farmington, vi–vii.1944, 2 & (Bohart) (USNM); Colorado: “Colo.”, coll. Coquillett, & (USNM). Dyscrasis hendeli: Type material: Holotype (: “Dallas Tex. / III 29.07”, “FCBishopp / collector”, “Dyscrasis / hendeli / n. sp.”, “Type / No. 43575 / U.S.N.M.”, allotype &, labels as in the holotype; 14 of 17 paratypes mentioned in original description: Dallas, Texas, 17.iii.1908, &, idem, 20.iii.1908, (, &, idem, 29.iii.1908, 2 (, idem, 30.v.08, & (Bishopp); Uvalde, Tex. in trap, 16.xi.1915, ( (Bishopp No. 5672); Matamoros, Mexico, in traps, 12.ix.1931, (, 3 & (Stephens); “Mercedes, Texas, in trap”, 26.viii.1931, ( (Heard); “San Benito, Tex. on office window, 26.v.1930, ( (Greer); “Tex.”, & (USNM). Non-type material: USA: Arizona: Baboquivari Mts., 23.ix.1923, (, & (Bryant) (SIZK); Cave Creek, 6.x.1951, ( (Sturtevant); Nogales, 31.vii.1944, 2 & (Sturtevant) (USNM); Mexico: Morelos: Cañon de Lobos, 17– 22.iii.1960, 2 & (Benschoter) (SIZK); “Morelos”, 11.1964, (, & (Maltbyr) (USNM). Myennis mandschurica: Type material: Paratype (: [China]: “Mandschurei, Charbin / 7.VII.1953, W. Alin”, “Myennis mandschurica m. P.T. det. M. Hering 1955”, “Paratypus” (DEI). Non-type material: Russia: Primorskiy kray: Tigrovaya, Suchan district, 2.vii.1927, & (Stackelberg) (ZISP). Myennis sibirica: Type material: Syntype &: [Far East Russia: Khabarovskiy Kray: Raddevka] “sibirica” [Portschinsky’s handwriting] (ZISP). Non-type material: Russia: Khabarovskiy Kray: Malyy Khingan, Dichun River, 1.vii.1980, 2 & (Shatalkin); Pashkovo, bleeding willow, 1.vii.1980, & (Ozerov); Primorskiy Kray, 40 km SE of Ussuriysk [Kamenushka], ( (Ozerov] (ZMUM). Myennis tricolor: Type material: Syntypes 5 (, 3 &: [Turkmenistan:] “Sary Yasy / III.87”, “Reitter 1894 / Turkmenien”, 1 ( and 1 & with red labels “Type” and 4 ( and 2 & with yellow labels “Paratype” (NHMW). Non-type material: Uzbekistan: Samarqand, 27.viii.1938, 8.vi.1939, 3 & (collector unknown) (SIZK); Buhara, 10.vi.1928, & (Zimin) (HNHM); Daynau [= Denow], [ex] turanga [Populus euphratica], 13.iv.1973–13.v.1973, 2 ( [Krivosheina(?)] (HNHM); Tajikistan: “Stalinabad” [= Dushanbe], 23.VIII.1942, & (Gussakovski) (ZISP). Neodyscrasis steyskali: Non-type material: Mexico: “Iguala, Finca Tlatocan” [Guerrero], 07.1987, & (Figuera) (USNM); Guatemala: “Guatemala”, 6.iv.1931, ( (Axel Pira); Guatemala City, 29.v, 4, 10, 13.vi.1931, 2 (, 4 & (Bates) (USNM); Venezuela: Distrito Federal, 14 km N Colonia Tovar, 1750 m, 21–25.i.1983, (, & (Flint) (USNM); idem, ( (SIZK). Oedopa ascriptiva: Type material: Syntypes: (: “S. Colorado 879 –I”, “coll. Hendel”, “Oedopa / det. Hendel \ ascriptiva / TYPE”; &: “Morrison / Colorado”, “coll. Hendel”, “Oedopa / det. Hendel \ ascriptiva H.” (NHMW). Non-type material: “Col.”, &; “TDA Cockerel / Co. Col”, &; “Colorado, Rio Grande Co. 8000', South Fork, 20.vi.1982, & (Wirth) (USNM); Echo L, 600', Mt. Evans, Colo. 17.vii.1961, ( (Mason) (CNC). Oedopa capito: Type material: Syntypes: 4 (, 1 &: “Neb.”, “capito” and “capito m.” [Loew’s handwriting on 2 specimens], “Coll. Loew”, “Type 13260” (red label) (MCZ; CNC). Non-type material: Canada: Alberta: Lethbridge, 3.vii.1923, ( (Gray) (CNC); California: Palm Springs, on Chilopsis, 3.v.1955, ( (Mason) (CNC). USA: Mississippi: Lafayette Co. (, & (Hull) (CNC); Nevada: Clark 36º05.2' N, 114º58.8′W, 10–11.v.2001, & (D. and W.N. Mathis) (USNM). Oedopa sp. near ascriptiva: Canada: British Columbia: Osoyoos, 20.vii.1966, ( (Lazorko) (SIZK); Oliver, 10 (, 1 & (MacDougall and Kelton) (CNC); Alberta: Lethbridge, 3.vii.1923, ( (Gray) (CNC); Saskatchewan: Redfield, 10.vi.1978, ( (Lazorko) (SIZK). USA: Utah:

575 BIOTAXONOMY OF TEPHRITOIDEA

Strawberry Valley, Wasatch Co. dry river bed and bank, 9.vii.1961, & (Chilcott); Colorado: Great Sand Dunes Nat. Mon. 7600', 24.vii.1968, ( (Becker) (CNC); New Mexico: Highrolls, 11.vi.1902, ( (“capito Lw. det. Cresson”) (HNHM). Paroedopa punctigera: Type material: Syntypes: &: “Catal Spgs / 29.4 Ar”, “Coll. Hubbard / and Schwarz”; “Type / No 4471 / U.S.N.M.” less than red papermore than “Paroedopa / punctigera Coq.”; &: “Ckll. 1457 / StaFe July” [not marked as a type] (USNM). Non-type: USA: Arizona: Upper Sabina Cyn., Tucson, Santa Catalina Mts., 1, 24.v. 1971, 3 (, 2 &; Redington pass, 5 mi. E Tucson, 18.x.1970, (; Cochise, Strong-hold, 12.iii.1971, ( (Foote) (CMP); Cochise Co., Huachuca Mts., 5354 Ash C[an]y[o]n. Rd., 0.5 mi W of H[igh]w[a]y 92 / E, 5100 ft, at UV light, 3.vi.1992, & (McFarland), S[outh]W[est] Res[earch] Sta[tion], 5 mi SW Portal, 5400 ft, 06.1967, ( (USNM); New Mexico: Alamogordo, 10.iv–1.v.1902, 2 (, 1 & (NHMW); idem, 19.iv.1902, (; idem, 26.iv.1902, & (Cresson det.) (HNHM ); Las Vegas, “9.8”, & (H. S. Barber) (Cresson det.) (NHMW); idem, 6.v.1902, &; Carlsbad, 3.v.1905, ( ([?]e) (USNM); Texas: Big Bend N.P., Chihuahuan desert nr. Nugent Mt., 6.vi.1967, ( (dissected) (Blanchard) (SIZK). Pseudodyscrasis scutellaris: Non-type material: Mexico: Tlaxcala: Calpulalpan, “Collectado en Maguay”, 25.v.1967, & (Martell); México: Chapingo, 26.viii.1962, ( (Pacheco) (USNM). Pseudotephritina cribellum: Type material: Syntype ( of S. cribellum: [USA]: “Neb.” [printed], “cribellum / m.” [Loew’s handwriting on green paper bar], “Loew/Coll.”, “Type / 13259” (MCZ); holotype & of S. cribrum: “cribrum / m.” [Loew’s handwriting on green paper bar], “Loew/Coll.”, “Type / 13258” (MCZ). Non-type material (only labels for new records are listed): Iowa: Ames State Nursery, Ames, 25–30.vi, 2, 13.vii.1972, Malaise trap, (, 4 & (Pinger) (USNM); Michigan, Monroe, 29.vii.1958, &; idem, 20.ix.1958, & (Steyskal) (HNHM); Ohio: 7 mi E Kent, Ohio, 24.vi.1963, & (Suman) (USNM); South Dakota: Yankton, 30.vi.1960, (, 3 & (Edman); Big Stone Cy., 14.vi.1932 (Aldrich) (USNM); Wyoming: Devils Tower Nation. Monument, 19.viii.1969, 2 ( (B. Foote) (CMP). Pseudotephritina inaequalis: Type material: Holotype &: “De Beque Colo. / July 29 1922”, “ERKalmbach / collector”, “Pseudotephritis (Pseudotephritina) / inaequalis / type / JRMalloch det.”, “Type / No. 43457 / U.S.N.M.” [red paper label] (USNM). Non-type material: Canada: Alberta: Lethbridge and Medicine Hat, 25.viii.1922, 4.vii.1927, 10.vii.1937, 6 (, 4 & (Gray; Seamans; Bryant) (CNC; USNM); USA: Utah: Emery Co., Wild Horse Creek 1.5 mi W Wild Horse Butte, swept from leaves of cottonwood (Populus), 26–27, 31.vii.1982, 2 & (Menke) (USNM); over 50 specimens from most localities listed by Steyskal (1962) were reexamined. Pseudotephritis approximata: Non-type material: USA: District Columbia: Washington, 16.viii.1913, & (USNM); Indiana: La Fayette, 27.vi.1916, & (Aldrich) (USNM); Iowa: Ames, State Nursery, Malaise trap, 28, 30.vi.1972, 2 (, & (Pinger) (USNM); Kansas: Lawrence, 1.06, 2 &, 13.vi.1900, ( (Kahl) (CMP); Maryland: Colesville, 17.vi.1975, (, 4.vii.1976, & (Wirth) (USNM); Mississippi: Oxford, 05.1945, 3 & (USNM); Missouri: Columbia, Malaise trap, 7 am-4 pm, 9.vii.1961, & (Parker) (USNM); Nebraska: Otoe Co., Dunbar, 19.vii.1969, & (Wirth) (USNM); New York: Fleetwood, “28–30 July”, & (Bromley coll.) (USNM); Oklahoma: Sequoyah Co., Tenkiller Lake, 3 mi W Blackgum, 11–14.vii.1979, ( (Davis) (USNM); Pennsylvania: “24 Jul.”, 7 (, 4 &; Pittsburgh, Shealy Park, 29.vii.1908, 3 &; Hershey, “4-7- 90”, & (Yackley) (CMP); Virginia: Falls Church, 30.vii.1960, (, & (Wirth) (USNM); West Virginia: Hardy Co., Lost River State Park, 11.vii.1974, 2 (, 2 & (Steyskal) (USNM); about 15 more specimens from various localities in Virginia, Maryland and Pennsylvania.

576 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Pseudotephritis corticalis: Type material: Syntypes Stictocephala corticalis Loew: (: “7653” [black ink; paper bar w/ 2 red lines], “29.” [printed on blue paper], “corticalis / Fitch [?] Mspt.” [Loew’s h/w; second word not understood], “Loew/Coll”, “Type / 15082 [red square]”; &: “corticalis” [Osten Sacken’s [?] h/w], “Loew/Coll”; “Type / 15082 [red square]” (MCZ); Syntype Myennis trypetoptera Hennig: &: “vic. Luga NW Russia / Tolmatschevo 10.VI.1937 / Stackelberg”, “Micropraep. Fluegel”, “Typus” (DEI). Non-type material: Canada: Alberta: McMurray, 14, 17.vii. 1953, 4 (, & (W. J. Brown) (CNC); British Columbia, Chilcotin, 18.vi.1920, ( (Buckell); Vernon, 8.vi.1936, (, & (Leech) (CNC); Manitoba: Whitewater Lake, 4 mi N Whitewater, 22.vi.1958, ( (Hurley) (CNC); Ontario: Gibson Lake, 6 mi E Go Home Bay, 6.v.1959, ( (Chilcott) (CNC); Quebec: Beechgrow, 45º39’N, 76º08’W, 16.v.1962, 20.v.1970, 2 & (Vockeroth) (CNC); Saskatchewan: Prince Albert, 31.v.1948, ( (Vockeroth) (CNC); USA: Ohio: “W’[ater]loo. T[o]wn[shi]p”, Athens Co., 16.xi.1937, sex? (Stehr); Utah: Roosevelt, 19.vii.1950, & (Harmston); about 60 specimens from most localities listed by Steyskal (1962) (USNM); Russia: East Siberia: Amur Oblast: Zeya, 17.vi.1977, ( (Belov) (ZMUM); Klimoutsi, 40 km W of Svobodnyy, 3.viii.1958, (, & (G. Zinovyev) (HNHM); Primorskyy Kray: Vinogradovka, 30.v.1929, & (Kiritshenko) (ZISP); Yakovlevka, Spassk Distr., 6.viii.1926, ( (Dyakonov and Filipyev) (HNHM); Ternei, 17.v.1936, 2 & (Grunin) (ZMUM); Mongolia: Central Aimak: Nucht in Bogdo ul, 12 km SE of Centre 1650 m, 3.vi.1967, & (Dr. Z. Kaszab exped.) (HNHM). Pseudotephritis metzi: Holotype &: “Cold Spring / Harbor L.I.”, “Type 7718” [red paper], “HOLOTYPE / No. metzi” (MCZ) [red paper]. Pseudotephritis millepunctata: Type material: Holotype &: “St. Siza, Sutschan, Ussuri- Gebiet”, 15.vi.1927 (Stackelberg) (ZISP). Non-type material: China: “Han-tao-he-tzu, Manchukuo”, 20.viii.1936, ( (Weymarn) (CMP); Heilongjiang: “Gaolinzsy”, 2–8.vii.1939, & (Alin) (DEI). Russia: East Siberia: Chita oblast: Undurga River, 9.vii.1977, ( (Kovalev) (ZMUM); Far East Russia: Amur Oblast, 40 km W of Svobodnyy, 3.v.1958, & (G. Zinoviev) (ZISP); Pashkovo, on bleeding willow, 16.vii.1980, & (Ozerov) (ZMUM); Khabarovsk Kray: Jewish Autonomous Oblast: Malyy Khingan Range, Dichun River, 15–16.vii.1979, 2 ( (Shatalkin) (ZMUM); Primorskyy Kray: Kamenushka 30 km SE of Ussuriysk, 21.viii.1980, 2 & (Ozerov) (ZMUM). Pseudotephritis vau: Non-type material: Canada: Alberta; British Columbia; Northwest Territories: Norman Wells, 16.viii.1969, & (Shewell) (CNC); Nova Scotia: Smiley Brook, nr. Brooklyn Hants, 18.vii.1968, ( (Ferguson) (USNM); Ontario, Quebec, USA: Alabama: Kushla, 18–31.x.1916, 3 (, & (Sturtevant); Connecticut: Redding, 5.vi.1932, & (Melander); Shelton, 10.viii.1932, &; New Haven, 9.v.1916, ( (Plumb); Milford, 9.viii.1924, & (Kaston); Georgia: Atlanta, 21.vii.1947, & (Fattig); Florida: Florida Caverns Sta. Park, Jackson Co., 26.v.1973, 2 & (Wirth); Gainesville, 4.x.1981, & (Gross) (USNM); Iowa: Ames, 15.v.1970, 6 & (Pinger); Kansas: Oswego, Labette Co., 9.viii.1966, 2 & (Hevel); Louisiana: Buras, 3 (, 2 & (Heatrick); Maine: Cherryfield, 28.vii.1929, &; Massachusetts: Springfield, 13.6.1901, (; Woods Hole, 6.vi–19.vii.1923, 7 (, 2 & (Sturtevant); Oklahoma: Sequoyah Co., Tenkiller Lake, 3 mi W Blackgum, 6–9.vii.1979, (, & (Davis); Oregon: Benton Co., McDonald Forest, Oak Creek, 30.vi.1971, ( (Steyskal); Pennsylvania: Phyladelphia, 6.05, 2 ( (HNHM); West Virginia: Kanawha Co., S Charleston, ex white oak, 4.vi.1982 , & (Adler); altogether about 50 (, 90 &, also from localities cited by Steyskal (1962) and Canadian provinces listed above. Stictomyia longicornis Bigot: Non-type material: USA: Arizona: Slobe, 24.iv.1938, (, 2 &

577 BIOTAXONOMY OF TEPHRITOIDEA

(F.H. Parker) (SIZK); 2 mi E of Yarnell, Yavapai Co. 27.vi.1966, 2 (, 2 & (Westcott); Catalina Mts. 8.v.1944, (, 2 & (Frost, Jr.) (USNM); New Mexico: Alamogordo, 6.vi.1902, ( (Cresson det.) (NHMW); idem, 10.v.1902, & (USNM); Texas: S. Antonio, on Opuntia, 31.05, (, & (Morgan) (SIZK); Mexico: Ciudad Victoria, 09.1965, (, 6 & (Krauss) (USNM); “Mexico, 1883”, 6 (, 3 &, “445”, “Opuntia”, (, “Mexico”, 3 (, “Mexico, Sta. Fe, 1871”, & (Bilimek) (NHMW); locality not given: & (NHMW). Stictomyia punctata Coquillett: Type material: Holotype &: “Foot of / Little Mtn / Mesilla / Valley”, “on red / cactus flower / Apr. 10. / (Ckll.) | Ckll. 5357”, “Stictomyia / punctata / Coq.”, “Type / No. 4472 / U.S.N.M.” (USNM). Non-type material: USA: Idaho: 5 mi NE Malta, 7.ix.1962, & (Barr); Arizona: Tucson, “Nichol. | sublot /656”, 27.vi.1930, &; idem, Pima Co. 4.vi.1952, (, & (Casier, Gertsch, Schrammel); idem, 5 ml NE Tucson, Pima Co., “Biol. Note 7113”, 4.iv.1971, & (Foote); Santa Rita Range Res. Pima Co., 4.vi.1957, swept from mesquite, 2 (, 3 & (Butler); S[outh]W[est] Res[earch] Sta[tion], 5 mi SW Portal, 5400 ft, 06.1967, ( (USNM); New Mexico: Alamogordo, 5.vi.1902 (USNM). Ulidiotites dakotana Steyskal: Type material: Holotype (: South Dakota: “Martin, S.D. / July 6–1924”, “H.”; “Holotype / Ulidiotites / dakotana / Steyskal” [red paper, Steyskal’s handwriting] (USNM). Paratypes: South Dakota: 3 (: “Martin, S.D. / July 6–1924 | H.”, “Paratype / Ulidiotites / dakotana / Steyskal [red paper, Steyskal’s handwriting] (USNM). Non- type material: USA: Wyoming: D. Johnston Pl. Station, Glenrock, “341”, 31.v.1974 (Lavigne) (new record); Nevada: Humboldt Co., 10 mi N of Winnemucca, 23.vi.1971, 21 (, 10 & (Oman) (new record); Utah: Coral Pink Sand Dunes, Cane Co. 3.vii.1966, 14 (, 4 & (Allen) (new record); Kansas: Medora, Sand Hills, 29.vii.1936 (Oman) (all USNM) (new record); Mississippi: Lafayette Co. & (Hull) (CNC) (new record).

APPENDIX 2

List of characters used in the cladistic analysis The character distributions among the genera are shown in Table 1. Head 1. Shape: 0) almost as high as long and at most 1.5 times as wide as long; 1) 1.8–2.2 times higher and wider than long. 2. Face between antennae: 0) without a spot; 1) with a velvety-black spot; 2) with a transverse bar. Unordered. 3. Gena: 0) less than half as high as eye; 1) more than half as high as eye. 4. First flagellomere in lateral view: 0) short oval or round; 1) 3-3.5 times as long as wide; 2) more than 4 times as long as wide. Thorax 5. Dorsocentral setae: 0) two or one postsutural; 1) more than 2 postsuturals; 2) more than 2 postsuturals and 1 presutural setae. Pseudodyscrasis scutellaris, Neodyscrasis steyskali and Dyscrasis hendeli differ from other Myennidini in having additional setae, including more than 2 dorsocentrals, which is an apomorphic state, compared to the chaetotaxy of other Myennidini; D. hendeli, moreover, has presutural dorsocentrals. 6. Supra-alar setae: 0) one; 1) two. 7. Postalar area: 0) without velvety black spot; 1) with velvety black spot.

578 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

A similar spot occurs in Pseudotephritina, Dyscrasis, Neodyscrasis and Pseudodyscrasis, but is absent in other Myennidini. 8. Intra-alar area (in female): 0) with single strong intra-alar seta; 1) with cluster of setulae in addition to weak intra-alar seta. 9. Scutellum: 0) completely microtrichose; 1) largely shiny or at least with 2 large shiny spots. Legs 10. Midtibial pattern: 0) uniformly yellow; 1) with black ring on apical half and completely darkened basal half; 2) with 2–3 dark rings and yellow ring or wider area in basal third. 11. Thickened apicoventral setae of midtibia: 0) one of the setae is twice or more as long as others; 1) three or more setae subequal, the shortest 0.6–0.7 times as long as the longest. Wing 12. Basal two thirds of costal cell: 0) with a hyaline spot crossing cell and separating basal brown spot from apex; 1) with hyaline incision at anterior margin; 2) completely brown. Ordered.

13. Vein R1: 0) apically setulose; 1) bare.

14. Medial spot in cell r1: 0) not extending posterior to vein R4+5; 1) extending posterior to vein

R4+5.

15. Apical spot on vein R4+5: 0) separated from spot at apex of cell r2+3; 1) fused with spot at apex

of cell r2+3. 16. Brown area bordering crossvein R-M: 0) without hyaline marks; 1) with hyaline spots on each side. 17. Distance between crossveins R-M and DM-Cu: 0) as long as R-M; 1) shorter than R-M. 18. Crossvein DM-Cu: 0) straight; 1) sinuous. 19. Stump vein of M in discal cell: 0) absent; 1) present. 20. Vein M: 0) not bent anteriorly; 1) conspicuously bent, ending before wing apex.

21. Extension of cell bcu: 0) as long as or shorter than vein A1+Cu2; 1) longer than A1+Cu2. 22. Second crossband (from costal cell through middle or base of discomedial cell): 0) ending

on medial portion of vein A1; 1) ending apical to apex of A1+Cu2.

23. Crossband extending from costal cell to A1+Cu2: 0) crossing middle of cell dm; 1) crossing base of dm. Abdomen 24. Abdominal tergite 3: 0) uniformly microtrichose; 1) with darker spots at seta bases. 25. Female sternite 4: 0) subquadrate; 1) elongate oval. 26. Setulae on pleural membrane of abdomen: 0) absent; 1) present. 27. Metaphallic plate: 0) absent; 1) present. 28. Metaphallic plate: 0) apically pointed; 1) truncated or incised. 29. Metaphallic plate length: 0) moderate or short; 1) long and narrow. 30. Number of prensisetae: 0) two; 1) three; 2) four to five. 31. Phallus: 0) bare; 1) finely trichose; 2) long spinulose, at least on apical third. Ordered. 32. Neck of spermatheca (normally inverted inside spermatheca): 0) absent; 1) short; 2) long.

579 BIOTAXONOMY OF TEPHRITOIDEA

APPENDIX 3 Distribution Maps

Fig. 125. Distribution map of Pseudotephritis vau.

580 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 126. Distribution map of Pseudotephritis corticalis and P. approximata in the Nearctic Region.

581 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 127. Distribution map of Pseudotephritis corticalis and P. millepunctata in the Palaearctic Region.

Fig. 128. Distribution map of Callopistromyia.

582 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 129. Distribution map of Oedopa and Ulidiotites.

Fig. 130. Distribution map of Paroedopa and Stictomyia.

583 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 131. Distribution map of Myennis octopunctata in the Western Palaearctic Region.

Fig. 132. Distribution map of Myennis in the Eastern Palaearctic Region.

584 Kameneva and Korneyev / Myennidini, new tribe (Ulidiidae)

Fig. 133. Distribution map of Stictoedopa.

585 BIOTAXONOMY OF TEPHRITOIDEA

Fig. 134. Distribution map of Pseudotephritina.

Fig. 135. Distribution map of Dyscrasis, Neodyscrasis, n. gen. and Pseudodyscrasis.

586