01_Kameneva-1.indd UDC 595.773.4 IDENTITY OF SPECIES ASSIGNED TO THE GENUS CEPHALIA (DIPTERA, TEPHRITOIDEA) E. P. Kameneva1,2,* & V. A. Korneyev1,2 1Schmalhausen Institute of Zoology NAS of Ukraine, vul. B. Khmelnytskogo, 15, Kyiv, 01054 Ukraine 2Museum für Naturkunde Leibnitz, Invalidenstr. 43, 10115 Berlin, Germany *Corresponding author E-mail: kameneva.elena@gmail.com E. P. Kameneva https://orcid.org/0000-0002-1568-5183 V. A. Korneyev https://orcid.org/0000-0001-9631-1038 urn:lsid:zoobank.org:pub:DBEC9AA4-4B25-4036-8269-7C41F6873394 Identity of Species Assigned to the Genus Cephalia (Diptera, Tephritoidea).  Kameneva, E. P., Korneyev, V. A. — Th e taxonomy of the genus Cephalia Meigen, 1826 has hitherto been confused. Th is paper comprises a review of the 13 nominal species assigned to the genus Cephalia to date. Of these, C. nigripes Meigen, 1826 has previously been considered a junior synonym of C. rufi pes Meigen, 1826, which is here found to be apparently the only species which actually belongs in the genus Cephalia. Regarding the position of the other species: Cephalia fascipennis Wiedemann, 1830, C. femoralis Wiedemann, 1830 and C. marginata Wiedemann, 1830 was already transferred to Setellia (Richardiidae) and C. caloptera Bigot, 1886 synonymized with Euphranta connexa (Tephritidae); C. bicolor Bigot, 1886 synonymized with Elassogaster linearis (Walker, 1849) (Platystomatidae); and C. quadripunctata Gimmerthal, 1842 tentatively placed in the genus Sapromyza (Lauxaniidae). Cephalia myrmecoides Loew, 1860 and C. fenestrata Coquillett, 1900 have been transferred to the monotypic genera Myrmecothea Hendel and Myiomyrmica Steyskal respectively, forming a clearly monophyletic lineage together with C. rufi pes within the tribe Cephaliini (Ulidiidae). Th e following synonymies are also established here for the remaining species: Acrosticta fulvicornis (Bigot, 1886) comb. n., = Cephalia fulvicornis Bigot, 1886 (Acrosticta fulvipes Coqullett, 1900 is possibly a junior synonym) (Ulidiidae); Herina fl avoscutellata (Becker, 1900), comb. n. (Ulidiidae), = Cephalia fl avoscutellata Becker, 1900; Rivellia maculipennis (Bigot, 1886) comb. n., = Cephalia maculipennis Bigot, 1886 = Rivellia brevifasciata Johnson, 1900 syn. n. (Platystomatidae). Th e taxonomic position of Cephalia within the subfamily Otitinae is discussed. K e y w o r d s : Diptera, Ulidiidae, Richardiidae, Platystomatidae, Otitinae, Cephaliini, taxonomy, nomenclature. Zoodiversity, 56(6): 447–462, 2022 DOI 10.15407/zoo2022.06.447 448 E. P. Kameneva,V. A. Korneyev Introduction Th e family-group taxon Cephaliinae was originally established by Schiner (1864: 63) for the single genus Cephalia Meigen, 1826 as a part of a larger family-group taxon, the Sepsinae. It originally included four nominal species from Europe: C. nigripes Meigen, 1826, C. rufi pes Meigen, 1826, C. formicaria Robineau-Desvoidy, 1830 (as a synonym of C. rufi pes) and C. quadripunctata Gimmerthal, 1842. Subsequently, due to its aberrant, elongated body and wing shape, Cephalia was associated with various superfi cially similar genera such as Tanypeza Fallén, 1820 (Tanypezidae) or placed in the family/subfamily groups “Ortalidae” (Becker, 1905) or “Platystominae” (Hendel, 1914) within the separate tribes Cephalina or Myrmecomyiina. As the diff erences in the ground plan of male genitalia of Platystomatidae and Otitidae were demonstrated by Hennig (1939, 1945), the genus Cephalia was transferred into the family Otitidae or subfamily Otitinae (Steyskal, 1965; Soós, 1984) on the basis of having a long, spinulose phallus and the inner surstyli with more than two prensisetae, as in most other Otitinae. Becker (1905) suggested that the name Cephalia was possibly preoccupied by Panzer (1805) in the Hymenoptera. Coquillett (1910) and Hendel (1914) accepted this statement and used the junior synonym Myrmecomyia Robineau-Desvoidy, 1830 as a replacement name instead, along with the tribal name Myrmecomyiini and subfamilial name Myrmecomyiinae, a point of view which was subsequently followed by Séguy (1934). Hennig (1939), however, established that Cephaleia Panzer, 1805 is not a senior homonym of Cephalia Meigen, 1826 and considered Myrmecomyia to be the invalid junior synonym of the latter name once again. Under Myrmecomyia, Hendel (1914) listed all the nominal species previously placed in combination with Cephalia, as follows: Cephalia bicolor Bigot, 1886 (Ceylon), a junior synonym of Elassogaster sepsoides Walker [now Platystomatidae]; Cephalia caloptera Bigot, 1886, a junior synonym of Euphranta connexa Fabricius [now Tephritidae]; Cephalia fascipennis Wiedemann, 1830, a species of Setellia [now Richardiidae]; Cephalia femoralis Wiedemann, 1830, a species of Setellia [now Richardiidae]; Cephalia? fenestrata, Coquillett, 1900 (N. America); Cephalia fl avoscutellata Becker, 1900 (Siberia); Cephalia? fulvicornis Bigot, 1886 (California) (“not a Myrmecomyia species!”); Cephalia? maculipennis Bigot, 1886 (N. America.) (“not a Myrmecomyia species!”); Cephalia marginata Wiedemann, 1830, a species of Setellia [now Richardiidae]; Cephalia myrmecoides Loew, 1860, a species of Myrmecothea [now Ulidiidae: Otitinae: Cephaliini]; Cephalia quadripunctata Gimmerthal, 1842, a species of Sapromyza [now Lauxaniidae]; Cephalia rufi pes Meigen, 1826 (“Mittel- und Süd Europa? Nord-Amerika”) (with junior synonyms Cepha- lia nigripes Meigen, 1826 and Myrmecomyia myrmecoides Robineau-Desvoidy, 1830). As a result, at the beginning of our study the genus Cephalia included only two nominal species in the Palaearctic Region: Cephalia fl avoscutellata and C. rufi ceps (Soos, 1984), and four nominal species — Cephalia fl avoscutellata, C. fulvicornis, C. maculipennis, C. rufi pes — in North America (Steyskal, 1965). Prior to this, Steyskal (1961) had established a monotypic genus Myiomyrmica and transferred Cephalia fenestrata into it. All records of the above, except for C. rufi ceps in Europe, were based on only a few or single fi nds and, quite probably, on dubious identifi cations needing verifi cation of their taxonomic positions and statuses. Recently, Wallace (2021: 54) wrote: “Steyskal (1987) indicates four species [from the Nearctic Region]; two were synonymized into other genera since the publication of Th e Manual of Nearctic Diptera”. He provided no sources for the second statement concerning synonymy, however, and we are unable to fi nd any. In 2005, while visiting the University Museum Oxford, VAK examined and took pictures of the species described by J. M. F. Bigot as Cephalia and considered to lie under that name until now; later, in 2005 and 2008, he also examined species described by Wiedemann in the collection of the Zoological Museum, University of Copenhagen. While preparing a review of the New World Otitinae, EPK examined these photos and identifi ed their taxonomic position. Surprisingly, these names were found to be senior synonyms of already known species. At our request, the holotype of Cephalia fl avoscutellata was photographed at the collection of the Museum of Natural History, Helsinki, and published on the collection website; its taxonomic position and status is also clarifi ed in this paper. Material and methods Th e specimens examined in this work are deposited in the following collections: LNHM — Latvian National Natural History Museum, Riga (Latvia); MNHNP — Museum national d’Histoire naturelle, Paris (France); MNKB — Museum für Naturkunde, Berlin (Germany); MCZ — Museum of Comparative Zoology at Harvard University, Cambridge (U.S.A.); MZH — Finnish Museum of Natural History, Helsinki (Finland); NHMW — Naturhistorishes Museum Wien (Austria); SIZK — I. I. Schmalhausen Institute of Zoology, Kyiv (Ukraine); 449Identity of species assigned to Cephalia (Diptera) SMNF — Senckenberg Museum für Naturkunde Frankfurt-am-Main (Germany); TAUI — Steinhardt Museum of Natural History, Tel Aviv University (Israel); UMO — Oxford University Museum of Natural History (United Kingdom); USNM — U.S. National Museum of Natural History, Washington, D.C. (U.S.A.); ZMUK — Natural History Museum of Denmark, Copenhagen (Denmark). Results Identifi cation of the types of nominal species originally described by Bigot (1886) and Becker (1900) as Cephalia resulted in the conclusion that none of them belong to this genus in its current concept, and are synonyms, either junior (as was previously known for Cephalia caloptera, which was already known to be a synonym in Tephritidae, or C. bicolor, which is a synonym in Platystomatidae — see below) or senior, having priority over some names in Ulidiidae and Platystomatidae. Analysis of information on the other species assigned to Cephalia shows that this genus is apparently monotypic and restricted to Europe, whilst all other species assigned to the tribe Cephaliini by Kameneva & Korneyev (2006) occur in Americas from the U.S.A. to Argentina and Chile. Cephalia fascipennis Wiedemann, 1830 (fi g. 1) Valid name. Setellia fascipennis (Wiedemann, 1830) (Richardiidae). Cephalia fascipennis Wiedemann, 1830: 469. Setellia fascipennis; Hendel, 1911 a: 196, 1911 b: 17; Steyskal, 1968: 53.12. M a t e r i a l . Holotype {: “C. fascipennis / Wied / Brazils /Lund”, “Type” [red paper], “Mus. Westerm.” (ZMUK). R e m a r k s : Th is species was transferred to Setellia by Hendel (1911 a, b). Fig. 1. Setellia fascipennis (holotype { of Cephalia fascipennis Wiedemann, 1830) (Richardiidae), ZMUK (photos by V. Korneyev, 2008): a — habitus dorsally; b — labels; c — head and thorax, left ; d — same, dorsally; e — abdomen, dorsally; f — wing; g — abdomen, mid and hind legs, left . 450 E. P. Kameneva,V. A. Korneyev Cephalia femoralis Wiedemann, 1830 (fi g. 2) Valid name. Setellia femoralis (Wiedemann, 1830) (Richardiidae). Cephalia femoralis Wiedemann, 1830: 469. Setellia femoralis; Hendel, 1911 a: 195, 1911 b: 17; Steyskal, 1968: 53.12. M a t e r i a l . Holotype {: “C. femoralis / Wied / Brazils /Lund”, “Type” [red paper], “Mus. Westerm.” (ZMUK). R e m a r k s : Th is species was transferred to Setellia by Hendel (1911 a, b). Cephalia marginata Wiedemann, 1830 Valid name. Setellia marginata (Wiedemann, 1830) (Richardiidae). Cephalia marginata Wiedemann, 1830: 470. Setellia marginata: 470; Hendel, 1911 a: 195, 377, 1911 b: 17; Steyskal, 1968: 53.13 M a t e r i a l . Holotype }: “aus Brasilien” [Freiress], “im Frankfurter Museum” (not located, not found in SMNF). R e m a r k s : Th is species was transferred to Setellia by Hendel (1911 a, b). Cephalia bicolor Bigot, 1886 Valid name. Elassogaster linearis (Walker, 1849) (Platystomatidae). Sepsis linearis Walker, 1849: 998. Fig. 2. Setellia femoralis (holotype { of Cephalia femoralis Wiedemann, 1830) (Richardiidae), ZMUK (photos by V.  Korneyev, 2008): a  — habitus dorsally; b  — labels; c  — head, left ; d  — head, dorsally; e  — thorax and abdomen, left ; f — abdomen, dorsally; g — wing. 451Identity of species assigned to Cephalia (Diptera) Elassogaster linearis: Steyskal, 1977: 137. Dacus sepsoides Walker, 1861: 163. Elassogaster sepsoides: Hendel, 1914: 22; Steyskal, 1977: 137. Cephalia bicolor Bigot, 1886: 385. R e m a r k s : Th is nominal species was transferred to the genus Elassogaster by Hendel (1914) and into synonymy with E. linearis and E. sepsoides by Steyskal (1977). Cephalia caloptera Bigot, 1886 Valid name. Euphranta connexa (Fabricius, 1794) (Tephritidae). Cephalia caloptera Bigot, 1886; Mik, 1887: 159. Myrmecomyia coloptera: Becker, 1905: 102 (misspelling). Euphranta connexa: Hendel, 1927: 68; Foote, 1984: 89; Norrbom et al., 1999: 147. R e m a r k s : Th is nominal species described from the Eastern Pyrenees (France) was placed into syno-nymy with Euphranta connexa by Hendel (1927). Cephalia fenestrata Coquillett, 1900 Valid name. Myiomyrmica fenestrata (Coquillett, 1900) (Ulidiidae). Cephalia fenestrata Coquillett, 1900: 24 (Kansas). Myiomyrmica fenestrata: Steyskal, 1961: 404; 1965: 646; Wallace, 2021: 9, 14, 69. M a t e r i a l . Type. Holotype }: U.S.A.: “Onaga // Kansas”, [Baker leg.], “Cephalia // fenestrata // Coq.”, “Type // No. 4474 // U.S.N.M.” (USNM), (examined). R e m a r k s : Steyskal (1961) established a separate monotypic genus for this species in his key to the Nearctic genera of Platystomatidae and Otitidae, with a short diagnosis in a key couplet based on the frons being entirely shining and the wing narrow with a pale brownish disk and a whitish spot. Judging from Steyskal (1961: Fig. 5), it also diff ers by the male epandrium bearing trilobate surstyli lacking large prensisetae. It shares its long antenna, very wide, subtriangular palp, bulging upper part of the occiput and lack of proepisternal seta with Cephalia, Myrmecothea and Tritoxa, these synapomorphies apparently supporting monophyly of a lineage represented by these four genera. Cephalia fl avoscutellata Becker, 1900 (fi g. 3) Valid name. Herina fl avoscutellata (Becker, 1900), comb. n. (Ulidiidae). Cephalia fl avoscutellata Becker, 1900: 385; Steyskal, 1965: 643; Soós, 1984: 57; Wallace, 2021: 9, 13 (fi g. 4C), 54. Myrmecomya fl avoscutellata: Becker, 1905: 105. O r i g i n a l d e s c r i p t i o n : “Diese Gattung wird von Loew zu den Ortaliden gerechnet; s. Loew. North American Diptera I. 43. Die vorliegende Art ist weder mit rufi pes Mg., nig- ripes Mg. noch mit quadripunctata Gimmerth. identisch. Auch Cephalia myrmecoides Lw. Wien. Ent. Monatschr. 1860. 8. 3. 9 aus Nord-Amerika ist eine andere Art, deren Schwinger und Schildchen schwarz sind, bei der auch die Flügel an der Wurzel eine schwarze Zeich- nung haben. Verglichen sind auch noch folgende Arten: Wiedem. Aussereurop. Zweifl . II. 469. Cephalia femoralis, fascipennis u. marginata aus Brasilien. Bob. Desvoidy Myod. 721. 723. Polystodes ichneumoneus. Myrmecomyia formicaria, micropezoidea. Weibchen. Th orax glänzend schwarz, in der Gegend der Schulterbeulen dunkel rostroth. Schildchen gelb mit 2 Borsten. Kopf rostroth; Stirn verdunkelt, auf der Mitte eine etwas fl ache Längsrinne, die von dunkler Bestäubung etwas matt ist. Die Fühler haben ein braun bestäubtes drittes Fühler- glied, die beiden ersten Glieder sind gelb; Borste ziemlich lang, nackt. Das gewölbte Unter- gesicht glänzend rothbraun mit schmalen etwas weiss bereift en Wangen. Hinterrücken und Hinterleib glänzend schwarz, die zerstreute kurze schwarze Behaarung ist kaum zu erken- nen; Legeröhre glänzend schwarz. Schwinger gelb. Beine schwarz mit rothen Hüft gelenken, auch die äusserste Wurzel und Spitze der Schenkel und die Kniee sind rostgelb. Die Flügel 452 E. P. Kameneva,V. A. Korneyev haben an der Spitze einen grossen schwarzbraunen Flecken; er beginnt an der Spitze der Randzelle, deren äusserste Ecke auch noch braungefärbt ist und verläuft allmälig verblas- send hinter der Mündung der vierten Längsader; innere Begrenzung buchtig. 4½ mm. lang.” [E n g l i s h t r a n s l a t i o n : Th is genus is assigned by Loew to the ortalids; sensu Loew, North American Diptera I: 43. Th e present species is not identical either to rufi pes Mg., nigripes Mg. or to quadripunctata Gimmerth. Also Cephalia myrmecoides Lw. 1860 […] from North America is another species, haltere and scutellum of which are black, and wing also having black markings at the base. […] Female. Th orax shiny black, dark rust-red in the area of humeral lobes. Scutellum yellow with 2 setae. Head rusty red. Frons darkened, in the middle with somewhat shallow longitudinal groove, which is somewhat dull from dark pollination. Antennae with brown microtrichose third segment, the fi rst two segments yellow. Arista rather long, bare. Th e arched lower face is shiny red-brown with narrow cheeks that are dusted with white. Postnotum and abdomen shiny black, short black hairs scattered about barely visible; Oviscape shining black. Halter yellow. Legs black with red hip joints, also the extremities of the roots and tips of the thighs and the knees are rusty yellow. Wing with large black-brown spot at the tip; it begins at the tip of the marginal cell, the outermost corner of which is also coloured brown, and runs gradually fading behind the apex of the fourth longitudinal vein; inner boundary indented. 4.5 mm. long] R e m a r k s : Th is enigmatic species was previously known only from its original description, without any illustrations. Th rough the kindness of Jere Kahanpää and Pekka Malinen (ZMH) the type was photographed at our request, which made it possible to clarify its taxonomic position. Th is species is much smaller than Cephalia rufi pes (which has wing length about 7–8 mm, whereas in H. fl avoscutellata it is about 4.5–4.7 mm) and does not possess any synapomorphies of the Cephalia  + Myrmecothea  + Myiomyrmica  + Tritoxa lineage (see discussion above) such as long antennae, dorsally bulging occiput, absent proepisternal seta, or thorax low anteriorly and very high posteriorly, which are unique for Cephalia + Myrmecothea + Myiomyrmica. It therefore does not meet the diagnoses of those lineages and certainly does not belong to Cephalia. It shares its oval, densely microtrichose fl agellomere 1, head shape, wing venation and pattern with the species of Seioptera Kirby Fig. 3. Herina fl avoscutellata (holotype } of Cephalia fl avoscutellata Becker, 1900) (Ulidiidae), ZMH http:// id.luomus.fi /GV.8191 (photos by P. Malinen, 2022, CC BY 4.0, partly modifi ed): a  — habitus dorsally; b  — same, left ; c — labels; d — head, left ; d — arista; f — wing. 453Identity of species assigned to Cephalia (Diptera) and Pseudoseioptera Hennig of the Seiopterini but has a single katepisternal seta instead of two (the synapomorphy of Seiopterini) and has a short pterostigma, and thus does not belong to that tribe. It also strongly resembles Proteseia steyskali Hernández & Kameneva, 1998 (provisionally assigned to Cephaliini) because of these characters but diff ers by the short pterostigma and vein R1 setulose only at apical part (pterostigma long and vein R1 entirely setulose in Proteseia), as well as in the shorter face and narrower palp having no subtriangular appearance. It strongly resembles species of the genus Herina Robineau-Desvoidy (Otitini), to which it is here transferred, in its bulky thorax which is wide and subequally high in its anterior and posterior parts. It is similar to Herina oscillans (Meigen, 1826) in the oval fl agellomere 1, rather high clypeus, widened palp and wing pattern restricted to the darkened basicostal and costal cells and pterostigma, as well as having a dark apical spot and crossveins without dark spots. Herina oscillans occurs from England, Morocco and southern Sweden to Hungary and Cyprus, mainly in mountain meadows, and diff ers in being a much smaller species (wing length 2.7–3.5 mm) with a reddish frons (dark brown in H. fl avoscutellata), black scutellum (brownish yellow in H. fl avoscutellata), and less expressed wing pattern. Herina is a large and hyperdiverse, subcosmopolitan genus of the Otitini occurring throughout Asia (mostly in the mountains), Europe, and North America down to Mesoamerica (Morgulis et al., 2013). It includes all those species of the Otitini which do not possess the characters of Dorycera, Hiatus, Otites, Melieria and Ceroxys, and is apparently non-monophyletic but not readily separable into smaller genera. Kameneva (see: Morgulis et al. 2013) established several groups of species, however, and among these was the oscillans group which included Herina oscillans, H. parva (Loew, 1864), H. pseudoluctuosa Hennig, 1939, H. merzi Kameneva, 2007 and H. lazi Kameneva & Korneyev, 2012. Further study of the genital structures of H. fl avoscutellata, including the structure of the male surstylus and female spermatheca, is necessary to decide whether it also belongs to this species group. Th e records from North America (Canada: N.W.T.) as C. fl avoscutellata (see: Steyskal, 1965; Wallace, 2021: USNM 1396546) are very probably misidentifi ed; detailed comparison of the morphological characters and COI barcoding mtDNA sequences of specimens from North America and Siberia is needed to clarify if they belong to diff erent taxa. Cephalia fulvicornis Bigot, 1886 (fi g. 4 ) Valid name. Acrosticta fulvicornis (Bigot, 1886) comb. n. (Ulidiidae) Cephalia? fulvicornis Bigot, 1886: 385. Cephalia fulvicornis: Steyskal, 1965: 643 (as “unrecognized”); Poole & Gentili, 1996: 203. Acrosticta fulvipes Coqullett, 1900: 24; Steyskal, 1965: 650 (possible synonym). M a t e r i a l . Holotype } Cephalia fulvicornis: “C. Fulvicornis. }. / Californ. J. Bigot” [bottom label], “Type” [red-boarded circle]. O r i g i n a l d e s c r i p t i o n : “Castanea, obscure nitida. Capite et antennis, omnino, fulvis; palpis castaneis; halteribus albis; abdomine basi rufo; pedibus fulvis, tibiis pallide fuscanis, tarsis fuscis, præter posticos basi fulvo pictos; alis hyalinis, costa basi, late, usque ad partent trientem et macula magna apicali, nigris. D’un châtain foncé et luisant. La tête et les antennes entièrement fauves; palpes châ- tains; abdomen à base roussâtre; balanciers blancs; pieds fauves, tibias un peu brunâtres, principalement les antérieurs, tarses plus foncés, les postérieurs largement fauves à la base; ailes hyalines, une assez grande macule à l’extrémité, le bord externe, à la base et jusqu’à l’extrémité de la deuxième nervure longitudinale, largement, noirâtres. Californie. — 1 specim. Les trois espèces marquées du point ? n’étant représentées dans ma collection que par un seul spécimen, il ne m’a pas été possible de les étudier suffi samment pour pouvoir affi rmer leur identifi cation avec le genre Cephalia.” 454 E. P. Kameneva,V. A. Korneyev [E n g l i s h t r a n s l a t i o n : (from Latin) Brown, darkly shiny. Head and antennae entirely yellow; palps brownish; halteres white; base of abdomen rufous; femora yellow, tibiae pale brown, tarsi brown, hind tarsi also basally yellow; wings hyaline, base of costa widely until third section, as well as large apical spot, black. (From French) Dark and shiny brown. Head and antennae entirely rufous; palps chestnut; abdomen with a reddish base; halteres white; legs fawn, tibiae a little brownish, mainly the front ones, tarsi darker, the hind tarsi broadly fawn at the base; wings hyaline, a fairly large macula at the tip; costal edge from base to the end of the second longitudinal vein broadly blackish. California. — 1 specimen. Th e three species marked with the query mark are represented in my collection by only one specimen, and it was not possible for me to study them suffi ciently to be able to affi rm their identifi cation with the genus Cephalia.] R e m a r k s: Th e holotype of Cephalia fulvicornis is very similar to Acrosticta fulvipes Coqullett, 1900, another nominal species also described from California (Coqullett, 1900). We examined the holotype } (“Los Angel[es Co.] / Cal[ifornia]”, “Collection / Coquillett”, “Acrosticta / fulvipes / Coq.”, “Type / No. 4473 / U.S.N.M.”), which looks to share all the characters of the latter including the number and position of frontal pits at bases of setulae, wing pattern and venation, coloration of legs, etc., except in the face and frons (raised ridges among shining pits at bases of setulae) being densely whitish microtrichose, which is the diagnostic character of A. fulvipes, whereas the holotype of A. fulvicornis has them partly shining. Th is could be due to partial damage of the fi ne vestiture by moisture or aging, or even due to individual variation. Additional material is therefore needed to confi rm their synonymy. Fig. 4. Acrosticta fulvicornis (holotype } of Cephalia fulvicornis Bigot, 1886) (Ulidiidae), UMO (photos by V. Korneyev, 2006): a — specimen, with labels; b — habitus dorso-laterally; c — head, dorsally; d — same, left . 455Identity of species assigned to Cephalia (Diptera) Cephalia maculipennis Bigot, 1886 (fi g. 5) Valid name. Rivellia maculipennis (Bigot, 1886), comb. n. (Platystomatidae) Cephalia? maculipennis Bigot, 1886: 385. Cephalia maculipennis: Steyskal, 1965: 643 (as “unrecognized”); Poole & Gentili, 1996: 203. Rivellia brevifasciata Johnson, 1900: 326; Hendel, 1914a: 177; Cresson, 1924: 229; Namba, 1956: 54; Steyskal, 1965b: 643, syn. n. O r i g i n a l d e s c r i p t i o n o f Cephalia maculipennis: “Antennis basi fulvis (segmentum tertium abest); capite palpis que nigris; thorace scutelloque nigro nitido, tergo ænescente; abdomine nigro nitido metallescente; halteribus nigris; pedibus fulvis; alis hyalinis, venis fl avis, transversis, lineisque dudibus minimis, transversalibus et basi, extrinsecus, macula apicali, costa denique, anguste, fuscanis. Les deux premiers segments des antennes fauves (troisième manque); tête et palpes noires, orbites fi nement bordés de grisâtre; thorax d’un noir luisant ainsi que l’écusson, tergum d’une nuance bronzée obscure; balanciers noirs; abdomen d’un noir luisant avec des refl ets métalliques obscurs; pieds fauves; ailes hyalines, nervure costale, les deux trans- versaîes et doux petites lignes transverses situées extérieurement vers la base, enfi n, une macule de médiocre dimension, s’anastomosant en arrière avec la nuance qui borde la ner- vure costale, noirâtres. Amér. septentr. (Rockey-Mount.). — 1 spécim. — Long. 4 millim.” [E n g l i s h t r a n s l a t i o n : (from Latin) Base of antennae yellowish brown (third segment is absent); head and palps black; thorax and scutellum glossy black, the back bronzed; abdomen black and shiny metallic; halteres black; legs yellowish brown; wings hyaline, veins yellow, extended, with few transverse veins, the transverse ones and their bases extrinsic; with apical blotch; costa narrow all the way, darkened. (From French) Th e fi rst two segments of the antennae fawn (third missing); head and palps black, orbits fi nely edged grayish; thorax shiny black as well as the scutellum, dorsum with dark bronze sheen; halteres black; abdomen shiny black with dark metallic sheen; legs fawn; wings hyaline, costal vein, the two transverse veins and the soft small transverse veins located externally towards the base, fi nally, a blotch of poor size, joining backwards with the shade bordering the costal vein, blackish. North America (Rockey-Mount.). — 1 specim. — Length 4 mm.] O r i g i n a l d e s c r i p t i o n o f Rivellia brevifasciata: “{ }. Lower half of the face shining black, upper half opaque, front brown, opaque, vertex with three shining black spots, the central one surrounding the ocelli, frontal and facial orbits silvery; occiput black, antennae reddish, tip of third antennal segment dark brown. Th orax and abdomen with uniform dark green [sheen]. Legs entirely light yellow. Th e four crossbands on the wings are obsolete or greatly abbreviated, the fi rst and second costal cells are hyaline; the fi rst band consists only of a spot in the marginal cell: the second and third bands do not extend beyond the fourth Fig. 5. Rivellia maculipennis (holotype { of Cephalia maculipennis Bigot, 1886) (Platystomatidae), UMO (photos by V. Korneyev, 2006): a — specimen, with labels; b — habitus dorso-laterally. 456 E. P. Kameneva,V. A. Korneyev longitudinal vein; the fourth, or apical band, about the same as in R. fl avimana, but not connected with the third along the costal margin. Length 4 mm. Th is species is nearest related to R. fl avimana Loew, from which it is at once separated by the obsolete bands and hyaline costal cells. Atco, June 18, 1893. I have also two specimens from Dr. Garry deN. Hough, collected by Mr. G. R. Pilate at Tift on, Ga., June 6th.” D i s t r i b u t i o n . USA: Colorado (Bigot, 1886); Georgia, Illinois, Kansas, Maryland, Michigan, Mississipi, New Jersey, New York, North Carolina (Namba, 1956). R e m a r k s : Th e holotype of Cephalia maculipennis Bigot is clearly conspecifi c with the other specimens described by Johnson and redescribed by Namba. We therefore consider Cephalia maculipennis Bigot, 1886 and Rivellia brevifasciata Johnson, 1900 to be synonyms. Cephalia myrmecoides Loew, 1860 (fi g. 6) Valid name. Myrmecothea myrmecoides (Loew, 1860) (Ulidiidae). Cephalia myrmecoides Loew, 1860: 24 (Washington, D.C.). Myrmecothea myrmecoides: Hendel, 1910: 310, 1914: 16; Steyskal, 1961: 404; 1965: 646; Wallace, 2021: 9, 14, 69. M a t e r i a l . Type. Holotype {: “Type /13244”, “Loew. / coll.” (MCZ). Non-type. USA: Maryland, Chevy Chase, 30.05.1964 2 { (P. P. Babiy leg.) (MCZ); Virginia: “DixieLndg //5.30 Va.”, coll. Hendel”, 2 } (NHMW). R e m a r k s : Hendel (1910) established a separate genus based on a single autapomorphy which diff erentiates it from Cephalia: i. e. anal lobe and alula absent; most other characters are similar to Cephalia rufi pes. Should Cephalia be found to contain more than one species, Myrmecothea should be considered a derived terminal taxon rather than its sister taxon. Cephalia quadripunctata Gimmerthal, 1842 Valid name. Sapromyza quadripunctata (Gimmerthal, 1842): Hendel, 1914: 22; Hennig, 1939: 68 (Lauxaniidae). Cephalia quadripunctata Gimmerthal, 1843: 684. Myrmecomya quadripunctata: Becker, 1905: 102. M a t e r i a l . Holotype }; not located; possibly LNNHM; not examined. O r i g i n a l d e s c r i p t i o n : “Th oracæ griseo, abdomine testacea, punctis quatuor; pedibus testaceæ. Fig. 6. Myrmecothea myrmecoides (holotype { of Cephalia myrmecoides Loew, 1860) (Ulidiidae), MCZ (photos by V. Korneyev, 2001): a — habitus right view; b — labels; c — head; d, e — left and right wing, respectively. 457Identity of species assigned to Cephalia (Diptera) Kopf, Fühler und Mundtheile rothgelb; Augen mit einer feinen weissen Linie umzogen. Mittelleib aschgrau. Schildchen und Hinterleib rostgelb, auf dem 3ten und 4ten Leibringe je zwei kleine erhabene schwarze Puncte oder Wärzchen. Beine rostgelb, Schenkel aussen grau schillernd. Flügel etwas gelblich getrübt, besonders am Vorderrande. Stark 2"' Ein }”. [E n g l i s h t r a n s l a t i o n : (from Latin) Th orax grey, abdomen reddish yellow, four dots, legs reddish yellow] (From German) Head, antennae and mouthparts reddish yellow; eyes outlined with a fi ne white line. Th orax ash grey. Scutellum and abdomen rusty yellow, on the 3rd and 4th abdominal segments two small raised black spots or tubercles. Legs rusty yellow, legs iridescent gray on the outside. Wings somewhat yellowish clouded, especially at the anterior margin. Size 2 lines [4.23 mm]. One }]. Cephalia rufi pes Meigen, 1826 (fi gs 7–9) Valid name. Cephalia rufi pes Meigen, 1826 (Ulidiidae). Cephalia rufi pes Meigen, 1826: 294; Becker, 1902: 230, remarks on the absence of the types in boththe Paris and Vienna collections; Schiner, 1864: Austria (Prater, the park on Danube islands in Vienna); Hennig, 1939: France, Germany, Italy, Spain; Soós, 1957: Hungary; Rohaček, 2006: Czech Republic (Bohemia), Slovakia; van Aartsen & Beuk, 2002: Th e Netherlands; Kameneva, 2007: review of European material. Myrmecomyia rufi pes: Séguy, 1934 Cephalia nigripes Meigen, 1826: 294; Becker, 1902: 230; Séguy 1934: as “variation” of rufi pes. Aft er Williston also in North America, Virginia, to be confi rmed. M a t e r i a l . Type. Syntypes: Cephalia rufi ipes: 1 }, [Klug, Berlin], 1 }: [Austria, Megerle von Mühlfeld] (not located; not examined). Holotype (?) } Cephalia nigripes [Germany:] “Aachen”, “Alte Sammlung”, “Cephalia / nigripes / M. / v. 29 / 7 A” [paper square, ink handwriting] [Baumhauer] (NHMW); holotype (?) } Cephalia nigripes [Country unknown]: “meigen \ 2442 / 40” [paper circle], “Cephalia / nigripes”, “MNNH, Paris / ED2996”, [bottom labels:] “2247”, “Cephalia / rufi pes” [old paper rectangle, ink handwriting], “Muséum national d’Histoire naturelle, Paris (France), Collection: Insects - Diptera (ED) Specimen MNHN-ED-ED2966” (MNHNP). Non-type. Austria: “Austria / coll. Egger”, 1 {, 1 } “rufi pes // det. Schiner”; “Schin. 1869” }; “Alte Samm- lung”, 19 specimens (C. rufi pes det. Schiner and Hendel); “Coll. Hendel”, “rufi ceps Mg. det. F. Hendel \ Myrme- comyia”, 1 { [dissected]; “Coll. Hendel”, ”rufi ceps Mg. det. Hendel \ Neusiedler See, Juni”, 1 {, 1 } [dissected]; “Bgst”, “rufi ceps / det. Bergenst[amm].”, 1 } (NHMW); Wien, 08.1861, “coll. H. Loew”, 1 }; “Austria, Brauer”, “coll. H. Loew”, 1 } [specimen heavily damaged by dermestids] (MNKB); France: Rambuiett, 4.07.1900, 1 {, 2.07.1946, 1 } (RBINH); “Cephalia // rufi pes // Lyon”, “rufi pes // coll. Winthem”, 1 } (NHMW); Pyrenées- Orientales, 610 m, Can Baills, 10km SW Th uir, 42.34N/02.39E, 8.06.2007, 1 } (Merz) (MHNG). France/Spain (?): “Pyrenaei Keitel”, “6626”, 1 } (“rufi pes / Meig.”) [head missing] (ZMHB); Germany: Karlsruhe, 30.07.1972, 1 } (Stritt) (SMNK); Spain: Pr. Cadiz, Hozgarganta-Tal bei Jimena 200m, 17.07.1979, 1 } (Schacht) (ZSSM); Pr. Salamanca, Villar de Ciervo, Las Coronas, 18.06–8.07.1995, 1 } (Tschorsnig) (SMNS); Switzerland: “Basel Imhot / v. Roser Coll.”, 2 {, 1 } “Myrmecomyia rufi pes Mg. [det. Roser] (SMNS); Israel: Har Hermon: Birket Hakar, 22.06.1971, 1 {, 3 } (Freidberg) (TAUI; SIZK). Fig. 7. Cephalia rufi pes (possible holotype } of Cephalia nigripes Meigen, 1826) (Ulidiidae), MNHNP (photos by V. Korneyev, 2005): a — habitus dorsally and labels; b — same, postero-dorsally. 458 E. P. Kameneva,V. A. Korneyev Fig. 8. Cephalia, non-type { (a) and } (b–f) from Har Hermon (photos by V. Korneyev, 2012): a — habitus right; b — same, left ; c — head, left ; d — thorax, left ; e — same, dorsally; f — wing. D e s c r i p t i o n : Head (fi g. 8, c) including appendages, and thorax, scutellum and legs, all rusty red to reddish brown, widely shiny, except the frontal vitta dull in the middle of its length and orbits with narrow silvery microtrichose eye margins. Face straight in profi le and conspicuously produced anteriorly, its upper and lower part (epistome) not separated by any depression or suture; subgenae (paired lateral extensions of face ventral of genae and separated from them by a suture from vibrissal angle to anterior tentorial pit) wide triangular, as high as gena itself. Clypeus moderately high. Antennal groove moderately shallow. Anterior orbital seta hair-like, short; posterior orbital seta moderately long. Inner vertical seta strong, outer vertical seta 0.66× as long as inner vertical. Ocellar seta vestigial. Postocellar seta conspicuous, but short. Postocular setulae short, forming no regular row. Occipital setulae lateral of foramen spinulose, 3–4× as long as postocular setae; genal seta strong. Antenna with short scape and pedicel, usually reddish or brownish yellow, fl agellomere 1 about 4–5 times as long as wide, narrowed apically, brown to black, microtrichose; arista 3-segmented, yellow at base, apically dark brown, very short pubescent. Palp enlarged, crescentric or subtriangular, brown to black, short black setulose and grey microtrichose, twice as high as clypeus. Prementum high, subshining, setulose. Labellum moderately short. 459Identity of species assigned to Cephalia (Diptera) Th orax (fi gs 8, d–e), including scutellum, mostly shiny. Scutum medially shagreened, matt, fi nely silvery microtrichose between two rows of short dorsocentral setulae; two short brown to black submedian vittae separated by yellow or brown vitta between two rows of short acrostichal setulae almost reaching posteriorly the level of supra-alar setae. Prescutellar area widely matt and shagreened; acrostichal seta indistinct, hair-like; posterior dorso- central and intra-alar setae weak and short, almost indistinct; 0 postpronotal, 0 anterior supra-alar, 1 posterior supra-alar and 1 postalar seta. Scutellum with 2 pairs of setae (basal shorter than half of apical scutellar seta). Pleura subshining, notopleural triangle and katepisternum faintly silvery microtrichose. Proepisternal seta absent; 1 anepisternal and 1 katepisternal setae moderately strong. Legs (fi gs 8, a–b), long, reddish yellow to dark brown; tarsi, mid tibia and hind leg conspicuously darker; short black setulose; fore coxa whitish microtrichose; femora narrow; midtibia with 1 apicoventral seta. Wing (fi gs 8, a–b, f) glossy with faint brownish-yellow tinge. Basicostal cell and base of costal cell brown; pterostigma dark brown to black. Vein R1 setulose only in apical part. Veins R4+5 and M1 slightly divergent apically. Brown apical spot aligned to costa from apex of cell r2+3 to m1. Crossvein r-m at level of R1 apex. Vein CuA slightly sinuate, posteroapical lobe of cell cua along vein CuA+CuP very short but conspicuous. Alula narrow, but present. Upper calypter narrow, white ciliate. Abdomen subshining or shining black. Syntergite 1+2 with conspicuous constriction. Male postabdomen. Epandrium (fi g. 9, a) subglobose, with ventro-mesally directed surstylus bearing one moderately acute subapical prensiseta and group of 3–4 mesally directed prensisetae (“subcercal prensisetae”) at base of each surstylus. Hypandrium (fi g. 9, b) as in most Otitini: almost symmetrical, with wide and moderately deep phallus guide dorsally forming moderately developed phallapodeme and anteriorly attached to large pregonites; each of the latter bearing 7–9 trichoid sensilla (“setulae”); postgonites (gonostyli) developed as two button-like sclerites at each side of basiphallus and bearing 5 trichoid sensilla. Phallus (fi g. 9, d) directed to left side and coiled at rest in a membranous pouch at left side of postabdomen; basally with thicker and denser, and apically with longer, sparser and thinner spines (“acanths”); no glans: apex with gonopore, membranous and bare. Ejaculatory apodeme with moderately narrow “fan”sz, well expressed “shaft ” and relatively small “foot” (fi g. 9, c). Female with shortened but exposed abdominal tergite 6, moderately long oviscape as long as tergite 5, and non-modifi ed, rather wide aculeus with oval cercal unit bearing numerous trichoid sensilla. Th ree subspherical spermathecae with sparsely papillose surface. Body 8–9 mm, wings 7–8 mm long. D i s t r i b u t i o n . Middle and Southern Europe: from France and Spain to Slovakia and Austria; Israel (fi rst record). R e m a r k s . Kameneva (2007) has already noted that Meigen (1826) described this species based on females, one of which he received from Mr Klug from Berlin and the other from Dr Megerle von Mühlfeld as collected in Austria; these specimens have not been located in the MNHNP, MNKB or NHMW collections. Th e female in Meigen’s collection (MNHNP) placed under C. rufi pes (No. 2247) does not meet the original description of that species; it has the mesonotum uniformly black. Becker (1902) also noted that it has entirely black legs (as in “nigripes”), but nevertheless marked it as a C. nigripes type specimen and considered it to be a male (sic!), which is obviously an error. Th e holotype female of C. nigripes “caught by Mr Baumhauer in August at Lustberge near Aachen” (Meigen, 1826) is believed to be in the NHMW collection, but its label diff ers from the original data in the month of collecting; the } of “Cephalia nigripes” in MNHNP instead has no obvious geographic label indicating that it is from Aachen and can hardly be its 460 E. P. Kameneva,V. A. Korneyev holotype. Th e specimen from Lyon (Winthem’s coll., NHMW) is certainly a non-type specimen. Th e records from North America (U.S.A.: Arizona, New Mexico) as C. rufi pes (see: Steyskal, 1965; Wallace, 2021: USNM 1396541) are very probably misidentifi ed; detailed comparison of morphological characters and COI barcoding mtDNA sequences of specimens from North America and Siberia is needed in order to clarify whether they belong to diff erent taxa. Discussion Th e genus Cephalia appears to be either monotypic or containing at most two or three poorly recognized species; most species previously assigned belong elsewhere in the superfamily Tephritoidea. Its taxonomy was confused, and records in the literature and specimens in collections are rare, despite C. rufi pes being one of the most peculiar of European fl ies, with a large 7–8 mm long ant-like body and Sepsis-like appearance (as in the family Sepsidae). Kameneva & Korneyev (2006) included Cephalia as a type genus in the tribe Cephaliini Schiner, 1864, which they extended to include the genera 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; Tritoxa Loew, 1873, and more recently, also Xycores Kameneva & Korneyev, 2017 (Kameneva et al., 2017). Th e phylogenetic position of the Cephaliini within the subfamily Otitinae needs additional study: despite numerous autapomorphies, these genera are believed to form an in-group within the wider and very possibly non-monophyletic Otitini (and possibly Fig. 9. Cephalia, non-type {, NHMW (photos by V. Korneyev, 2017): a — epandrium, postero-ventrally; b — hypandrium, ventrally; c — ejaculatory apodeme; d — phallus, detached. 461Identity of species assigned to Cephalia (Diptera) within the large, probably also non-monophyletic genus Herina within the Otitini), or alternatively they may lie within hitherto unclassifi ed “genera incertae sedis, possibly related to Cephaliini” (Kameneva & Korneyev, 2006). All of these groups appear to be represented exclusively by species whose larvae, as far as is known, infest rotting or living underground parts of perennial herbaceous plants, possibly only monocots, and occur mainly in the spring or early summer in meadows, prairies or steppes, oft en in mountain areas. Delphinia picta and some species of Tritoxa are documented as pests of bulbs of various Amaryllidaceae (onions, etc.) and other related families; and Myrmecothea was recorded in association of rotting vegetation (Ferrar, 1987), which is believed to be also the mode of larval feeding of the closely related Cephalia. Th ese issues are to be further considered in detail elsewhere (Kameneva & Korneyev, in prep.). Th is work was initially started in 2021 as a part of the current monographic project “Fauna of Ukraine. Th e higher tephritoid dipterans (Diptera: Ulidiidae, Platystomatidae, Pyrgotidae, Tephritidae)” (SIZK: III- 50-21) and fi nalized a part of studies conducted by EPK in 2022 at MNKB funded by the Philip Schwartz Initiative and Museum für Naturkunde Berlin, with the support of Christoph Häuser, Rudolph Meier, Anja Blessing, Jenny Pohl and Sven Marotzke (MNKB). We appreciate the kindness of Jere Kahanpää and Pekka Malinen for photographing a type specimen from the collection of ZMH and Andrea Hastenpfl ug-Vesmanis for detailed information on the Wiedemann type specimens deposited in the collection of SMNF. We thank David Clements for improving English text. James C. Trager kindly advised on Latin translations. We thank two anonymous referees for their constructive criticism and important comments and corrections in the manuscript. References Aartsen, B. van, Beuk, P. 2002. Family Ulidiidae. In: Beuk, P. L. Th ., ed. Checklist of the Diptera of the Netherlands. KNNV Uitgeverij Utrecht, 238–239. Becker, Th . 1900. Beiträge zur Dipteren-Faunaa Sibiriens. Nordwest-Sibirische Dipteren gesammelt vom Prof. John Sahlberg aus Helsingfors im Jahre 1876, und vom Dr. E. Betroth aus Tammerfors im Jahre 1877 Be- arbeitet von Th . Becker in Liegnitz. Acta Societatis Scientiarum Fennicae, 26 (9), 1–66 + 2 pls. Becker, Th . 1902. 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