Search

A checklist is presented of all 986 extant species of Diptera known from the archipelago of the Maltese Islands situated in the central Mediterranean. Species considered to be alien to the Islands are listed with annotations in Appendix 1. The history of dipterology applicable to the islands is outlined and the three important historical published records by Zetterstedt, Rondani and Bezzi & de Stefani-Perez are listed in Appendices 2–4. Species names that are synonyms are indicated where these were used in published records for Malta more or less after the Catalogue of Palaearctic Diptera was published. Species we consider to have been misidentifications are also indicated with an annotation in most cases. Vermileo immaculatus Carles-Tolrá syn. n. & Cuesta-Segura and Vermileo balearicus Wheeler syn. n. are proposed junior synonyms of Vermileo vermileo (Linnaeus).

5 ABSTRACT This doctoral thesis is focused on flat-footed fly families Opetiidae, Platypezidae and genus Microsania (Insecta: Diptera). The thesis consists of general introduction to the systematics, taxonomy, diversity, zoogeography, phylogeny, and biology of the flat-footed flies. Systematics and taxonomy are summarized based on historical as well as recent literature. Each important taxon is shortly introduced and information about its taxonomy, morphology, diversity, distribution, and biology is provided. The next two chapters deal with fossil species and research on species diversity. The following part is a summary and plan for future research. Final chapter consists of 9 published peer-reviewed articles (5 in impacted international journals and 4 in journals without impact factor). The first article is a molecular phylogeny study of relationships between genera of flat- footed flies. The results show Platypezidae consisting of two well-supported clades, the first with the subfamilies Melanderomyiinae + Callomyiinae and the second formed by subfamily Platypezinae. Genus Microsania was resolved as a separate lineage distant from Platypezidae which clustered with Opetiidae as its sister group, both together forming a sister group to Platypezidae. The genus Agathomyia proved to be paraphyletic....

The first Opetiidae known from the Southern Hemisphere is described—Puyehuemyia chandleri, gen. nov., sp. nov.—based on a female specimen collected in Valdivian forest in the Province of Osorno, south Chile. The Palearctic species Opetia nigra Meigen was also studied, allowing detailed comparisons. Features of the antenna, mouthparts, wing, and terminalia allowed the issue of the position of the family within the Eremoneura to be revisited. The inclusion of Opetiidae in the Platypezoidea is corroborated, possibly in a clade also including Lonchopteridae and Phoridae. The 3-articled condition of the styluslike arista in Puyehuemyia corroborates the hypothesis that the 2-articled condition in Opetia is independently derived, as it is in the Empidoidea and many schizophorans. Puyehuemyia chandleri has female terminalia typical of parasitoid groups, as does Opetia, although their life history is not known. Described Platypezoidea Cretaceous amber fossils are reviewed, and Lonchopterites is considered to be sister to the crown group of Opetiidae. The presence of an Early Cretaceous biogeographical layer in the Valdivian forest, associated with plant and animals disjunct from New Zealand, and similar to the beech forests in the Northern Hemisphere, is discussed.

The molecular phylogeny of flat-footed flies is inferred from analysis of DNA sequence data from the five mitochondrial genes 12S, 16S, COI, COII and CytB, and the nuclear gene 28S and discussed with the recent systematics based on morphological features. The Bayesian inference, maximum likelihood and maximum parsimony analyses included 42 species of 18 genera, representing all four extant subfamilies (Microsaniinae, Melanderomyiinae, Callomyiinae and Platypezinae) and all known genera except one (Metaclythia). Representatives of the brachycerous taxa Lonchopteridae, Phoridae, Sciadocerinae (Phoridae) and Opetiidae are used as outgroups, and Lonchoptera was used to root the trees. Our results show Platypezidae consisting of two well-supported clades, the first with the subfamilies Melanderomyiinae + Callomyiinae and the second formed by subfamily Platypezinae. Genus Microsania was resolved as a separate lineage distant from Platypezidae which clustered with Opetiidae as its sister group, both together forming a sister group to Platypezidae. At the generic level, the genus Agathomyia proved not to be monophyletic in any of the analyses. The species Chydaeopeza tibialis is sister to Agathomyia sexmaculata, and consequently, the genus Chydaeopeza Shatalkin, 1992 is a new junior synonym of Agathomyia Verrall, 1901. Bifurcated setae on legs of adult Platypezidae are documented as a new synapomorphy of the family, exclusive of Microsania. Outstretched wings and only a small overlap of their surfaces at resting position are considered a new synapomorphy for the subfamily Platypezinae. Other phylogenetically important characters defining main clades are documented, and their relevance/ validity in phylogenetic studies is discussed. The current systematic concept of Platypezidae is discussed, and new phylogenetic hypotheses are proposed.

Puyehuemyia, new genus Type species, Puyehuemyia chandleri, sp. nov. DIAGNOSIS: Maxillary palpus small, 1-segmented, antennal pedicel widening to apex, first flagellomere slightly wider midway to apex, arista 3-articled. Only R 1 setose among wing veins, R 1 not curved anteriorly toward C at apex, C circumambient. Tibial spurs absent, hind tibia slightly sinuous. Scutum with only one pair of posterior dorsocentral setae, acrostical setae absent. Scutellum with four strong setae evenly spaced at discal margin. Female segments 5���8 telescoped, ending as a sclerotized, possibly parasitoid-type ovipositor, cercus probably absent. COMMENTS: A generic status for this Chilean species seems well justified. Not only is the antenna strikingly different from Opetia, with a 3-articled stylus, but also there are features in the wing venation of Puyehuemyia that differentiates it from the remaining species of the family���particularly, the shape of Sc and R1, and the shape of the basal sector of Rs., Published as part of Amorim, Dalton De Souza, Silva, Vera Cristina & Brown, Brian V., 2018, Puyehuemyia chandleri, gen. nov., sp. nov. (Diptera: Opetiidae): remnant of a Cretaceous biota in Chile, pp. 1-28 in American Museum Novitates 2018 (3892) on page 4, DOI: 10.1206/3892.1, http://zenodo.org/record/5369634

Puyehuemyia chandleri, new species Figures 1–3, 12A MATERIAL EXAMINED: Holotype, female, Chile, Osorno, Parque Nacional Puyehue, Termas Aguas Calientes, Sendero Rápido del Chanleufú, Malaise trap, Jan. 14–Feb. 3, 2017, D.S. Amorim and V. C. Silva, cols. (MNHN). DIAGNOSIS: As stated for the genus. FEMALE: Total body length, 2.5 mm (habitus, fig. 1A). Head (fig. 1B, C): Blackish brown, rounded, eyes reddish, large, dichoptic, dorsal and ventral ommatidia of same size, eye entirely covered with pale ommatrichia. Labella cream yellow; maxillary palpus apparently 1-segmented, small, brownish, apically blunt, no apical sensory pit. Entire frons, occiput, and gena densely covered with setulae. Ocellar triangle clearly defined, ocelli of equal size, well separated. Frons with three well-developed fronto-orbital setae evenly spaced, anterior fronto-orbital and inner vertical slightly lateroclinate, mid and posterior fronto-orbitals inclinate; pair of reclinate strong ocellars, postocellars small. Eyes well separated dorsally at vertex and closely approximated ventrally, frons strongly triangular from a dorsal view. Antenna inserted at dorsal third of head, face slender, setose on ventral third, antennae close together. Antennal scape small, about half length of pedicel, with few setulae along distal margin; scape widening to apex, about one-half length of first flagellomere, with setae around distal half; first flagellomere wider midway to apex, with some scattered small setae on basal half, not produced at junction with arista or stylus; arista clearly with three slender articles of about equal length, texture similar to first flagellomere, covered with microtrichia, lacking setation. Face slender, brown, bare mesally, sparse setae on ventral third of parafacialia. Gena narrow, setose, with some stronger setae in irregular row. Thorax (fig. 2A, B): Cervical sclerite brown. Scutum dark brown anteriorly, gradually turning ochreous posteriorly; pair of dark brown stripes over undifferentiated dorsocentrals; one strong prescutellar dorsocentrals, no differentiated acrostichals. Two postpronotal setae, one strong and one minute; three strong presutural intraalars in irregular row; two large and three small notopleural setae; one strong supraalar. Scutellum ochreous, with apical pair of stronger, upcurved setae and pair of smaller, anteromarginal scutellars. Thoracic pleural sclerites not strongly sclerotized, ochreous, some areas brown or light brown on prosternum and proepimeron, anteriorly and dorsoposteriorly on anepisternum, on ventral two-thirds of katepisternum, on dorsoanterior corner of meron, on metepisternum and on ventral margin of metepimeron, around posterior spiracle, mesially on mediotergite, and at contact between scutellum and mediotergite laterally (fig. 2A). Pleural sclerites entirely bare of setae, except ventral half of proepisternum. Anterior spiracle small, rounded, surrounded by membrane, placed dorsally to proepimeron. Prosternum V-shaped, bare, more sclerotized ventrally (fig. 2B). Propleural suture sigmoid, proepimeron entirely fused to katepisternum at posterior end. Anapleural suture depressed mesially. Anepisternum cleft dorsal opening bearing single elongated basalare. Short dorsoposterior projection of katepisternum over meron. Katepisternum shield and meron shield well developed (respectively over mid- and hind legs). Meron more or less rectangular, reaching level of posterior spiracle, suture separating meron from mesepimeron and from metepisternum well marked. Partial suture of separation of mesopleurotrochantim and meron present. Limits of laterotergite from mesepimeron anteriorly and from mediotergite posteriorly well marked. Halter elongate, greyish, except lighter area basally; entirely bare of macrotrichia. Metanotum slender, but well characterized. Legs delicate, coxae dark brown, densely setose anteriorly, remainder of legs brown with light brown areas. Tibial spurs absent on all legs, hind tibia distally with regular comb of longer setae. Tarsomeres gradually shorter, with no denticles or enlarged setae, no teeth on tarsal claws. Foretibia distally with inner, less well-sclerotized area covered with slightly stronger setae, and distal regular comb of elongate setae; mid- and hind tibiae with some more developed setae distally in more or less regular row. Wing (fig. 2C): Length, 2.5 mm. Membrane light fumose brown, slightly darker along anterior margin, but without maculae. All wing veins bare of macrotrichia except R 1 with setae above along distal three-fourths. Hu present, well developed, Sc complete, reaching C at basal third of wing; R 1 long, gradually approaching C, reaching margin at distal threefourths of wing; R 2+3 long, reaching C close to wing tip, R 5 ending at wing tip (fig. 3A). C clearly circumambient. Transverse vein r-m discrete, in basal fourth of wing; M 1+2 (including sector basal to r-m) slightly longer than medial fork; M 4 thicker and more sclerotized than other posterior veins. Cells bm and br closed, crossvein m-m+M 3 (formerly dm-Cu) absent, very base of M 4 broken, m-cu present, cell cua present, closed, CuA+CuP reaching wing margin (fig. 3B). Abdominal tergites and sternites light ochreous brown, darker along posterior margin (fig. 3C). No abdominal muscle plaques. Abdomen more or less flattened; segments 1–4 well developed, sternite 1 nearly unsclerotized, posterior border of sternites 2–3 with unsclerotized lunular area, tergites 1–4 sclerotized, with row of slightly more developed setae along posterior margin; segments 5–6 slender, weakly sclerotized, tergite 5 with row of setae along posterior margin, sternite 5 with scattered setae, segment 6 bare, segments 7–8 strongly modified to constitute well sclerotized, elongated ovipositor (fig. 3D). Cercus probably absent. MALE: Unknown. ETYMOLOGY: The generic epithet refers to the locality—Puyehue National Park, in southern Chile —where the holotype was collected. Puyehue is the name of one of the big lakes in Osorno Province. The park was created in 1941 just east of the lake. In the Mapudungum, or Mapuche language, hue means “place,” while puye is the common name of the galaxiine fish, Galaxias maculatus (Jenyns, 1842). The specific epithet honors Peter Chandler, who has a long and substantial contribution to dipterology in general and on platypezids and opetiids in particular, including his excellent review of the European Opetiidae and Platypezidae (Chandler, 2001). DISTRIBUTION: This species is so far known only from the type locality, lowlands of Puyehue National Park. The vegetation is part of the Subantarctic Valdivian phytogeographic district (Gajardo, 1994), also known as Valdivian Forest. It is dominated by Nothofagus dombeyi Mirb. Oerst., also having the characteristic Myrtaceae Luma apiculata (DC.) Burret, the conifer Podocarpus nubigenus Lindl. and other austral floral elements. BIOLOGY: Not much can be said about the biology of this fly. It was collected with a Malaise trap in the lowlands (440 m) of a temperate rain forest, in a patch of primary forest recovering from previous anthropic influence, but still with large Nothofagus trees and a good amount of rotting wood and woodland detritus (fig. 4). The general structure of the environment where P. chandleri was collected is similar to the reports for O. nigra in Europe (e.g., Roháček and Ševčík, 2011; Tkoč and Roháček, 2014) and for the Japanese species of the genus (Saigusa, 1963). It is interesting to note that O. nigra adults have been reared from rotten Fagus wood (Speight et al., 1990; Chandler, 2001) and the larvae are still unknown. The sclerotized, piercing ovipositor of the females in Opetia species and in P. chandleri may suggest a parasitoid biology for the larvae, even though some tephritoids have a piercing oviscapt used to oviposit in fruits or stems. If this is correct, it would explain why the larvae have not been found although quite a number of adults have been reared from different localities. COMMENTS: The spermathecae cannot be observed without dissection. In Opetia there is a single, unpigmented spermatheca (Sinclair and Cumming, 2006). It may be the case that the spermathecae is similar in P. chandleri. Additional specimens would allow dissection of specimens, clarifying several morphological features, including details of the maxillary palpus and ovipositor. There are some dimorphic features in Opetia that may well apply also to Puyehuemyia. Females of Opetia also have dichoptic eyes, with undifferentiated dorsal and ventral ommatidia, and the anal lobe is not developed in the wing. Male eyes of Opetia nigra (fig. 5A, C) are holoptic, the dorsal ommatidia are larger than the ventral ones, and the wing has a well-developed anal lobe.

Altogether 18 species of the families Opetiidae and Platypezidae are reported from Romania, based on newly studied material and previously published records. The following three species are recorded from Romania for the first time: Agathomyia vernalis Shatalkin, 1981, Callomyia saibhira Chandler, 1976, and Lindneromyia hungarica Chandler, 2001. The presented differential diagnosis and a detailed redescription of body and genitalia of the male of Callomyia saibhira are based on one specimen which is the first male found in Europe. Information about distribution and biology of all 18 Romanian species is provided as well as photographs of selected important species. Finally, a new checklist of all Romanian species is given.

The main features of antennal segments 2 and 3 seen in the higher Diptera are described, including many that are not or inadequately covered in available publications. The following terms are introduced or clarified: for segment 2 or the pedicel—annular ridge, caestus, chin, collar, conus, distal articular surface, encircling furrow, foramen of articulation, foraminal cusp, foraminal ring, pedicellar button, pedicellar cup, rim; for segment 3 or the postpedicel—basal foramen, basal hollow, basal stem, postpedicellar pouch, sacculus, scabrous tongue, sub-basal caecum; for the stylus or arista—stylar goblet. Particular attention is given to the occurrence and position of the pedicellar button. The button is the cuticular component of a chordotonal organ, which perhaps has the role of a baroreceptor. It is present in the majority of families of Diptera, and possibly was present in the ancestral dipteran. Some generalizations about antennal structure are made, and a diagram showing the main trends in antennal evolution in the Eremoneura is provided. The general form of the antenna shows a transition from approximate radial symmetry (e.g., in Empis, Microphor, and Opetia) through to superficial bilateral symmetry (in many taxa of Eumuscomorpha), though there is usually much asymmetry in detail. More detailed descriptions and illustrations are given for selected taxa of Cyclorrhapha. The phenomenon of an additional concealed segment-like structure between segments 2 and 3, found among the Chloropidae, Pyrgotidae, etc., and formed from the basally flexible conus, is described. Some antennal features of the Calyptratae suggest a relationship to the Tephritoidea. Critical comments are made with regard to the recently published phylogenetic association of the Ironomyiidae with the Phoridae and the Pallopteridae with the Neurochaetidae. In discussing relationships of some taxa, a few non-antennal features, some needing further study, are mentioned, e.g., variation in separation of abdominal tergites 1 and 2 in the Opetiidae and other lower cyclorrhaphous families; the presence of supplementary claw-like terminal tarsal processes in the Lonchopteridae; the apparent restriction of the presence of barbed macrotrichia to the Phoridae, among lower cyclorrhaphans; variation in structure of the prelabrum in the Pyrgotidae; the microstructure of the facial cuticle in the Syringogastridae as compared with that of other families; the calyptrate-like development of the squama in some tephritoid taxa; variation in the subscutellum in the Conopidae; a feature of the larval posterior spiracles diagnostic for Coelopidae.

The fauna of Opetiidae and Platypezidae (Diptera) in the Gemer region (Central Slovakia) A review of the fauna of flat-footed flies (Diptera: Opetiidae and Platypezidae) in the Gemer region (Slovakia) is presented. Based on previously published records and material examined 1 species of Opetiidae and 17 species of Platypezidae are treated, each with comments on its general distribution, biology and faunistics, and/or nature conservation importance. Nine species of Platypezidae are considered particularly significant for the area because of comprising taxa that are endangered, stenotopic or generally rare. Of the five species newly recorded from the Gemer area, one, viz. Agathomyia collini Verrall, 1901, is a new addition to the Slovak fauna and three, viz. Agathomyia falleni (Zetterstedt, 1819), Protoclythia rufa (Meigen, 1830) and Polyporivora picta (Meigen, 1830) represent second records from Slovakia, the last species being re-discovered in Slovakia after more than 140 years.

Cyclorrhaphan Diptera are an extremely successful clade of ecologically and phylogenenetically important flies. Despite their significance the relationships among lower cyclorrhaphans ('Aschiza') remain controversial in spite of several morphologically based phylogenetic analyses. We sequenced a 2.7-kb fragment of 28S rDNA for taxa representing all lower cyclorrhaphan families (except Ironomyiidae), four schizophoran families, and seven empidoid out-group taxa. Phylogenetic analysis of these data strongly supports a monophyletic Cyclorrhapha (including the enigmatic taxon Opetia nigra) that is divided into two clades - a well-supported Eumuscomorpha (Syrphidae + Pipunculidae + Schizophora), and a weakly-supported Platypezoidea (all non-Eumuscomorpha). Consequently, the former grouping known as Aschiza, which included syrphids and pipunculids, is not a valid monophyletic clade. Within Platypezoidea, most of our analyses place Lonchopteridae as sister group to Opetiidae, and strongly support the monophyly of Sciadoceridae + Phoridae. Among the Eumuscomorpha we do not recover the monophyly of Syrphoidea (Syrphidae + Pipunculidae). Instead, all analyses place Pipunculidae as the sister group to Schizophora. This novel finding has never been proposed based on morphological data and will require more data (both molecular and morphological) and taxa to confirm.

A checklist of the Opetiidae and Platypezidae (Diptera) recorded from Finland.

An account is given of the three Aschiza families of Diptera: one species of Lonchopteridae, one species of Opetiidae and four species of Pipunculidae that occur in Malta and which are all new records for this country, peer-reviewed

Chimeromyina Arillo & Grimaldi, gen. nov. Diagnosis. At present, defined only on the basis of characters that are symplesiomorphic with respect to Chimeromyia: Arista terminal, with 3 aristomeres; wing with crossveins r-m and bm-cu separated by distance approximately equal to length of r-m; cell dm present, large, with crossvein dm-cu; vein CuA 2, cell cu p and A 1 [+ CuA 2?] present; anal lobe and alula present. Type Species. Chimeromyina concilia, n.sp. Etymology. Derived from the name of the sister genus, Chimeromyia.

Chimeromyina concilia Arillo & Grimaldi, sp. n. (Fig. 13) Diagnosis. As for genus. Description. HEAD: Fairly broad, width more than 1.5 x width of notum; eyes large, bare, facets not differentiated. Ocelli on small tubercle, tubercle with 2 pairs long setae. Antenna with apical arista; 3 aristomeres present. Vibrissae not present. THORAX: 0.44 mm long, bristly, each side of notum with 2 long notopleural setae and supra-alar seta; 2 pairs of dorsocentrals, row of long setulae anterior to dorsocentrals. Acrostichal setulae in one, extensive row, extended to posterior margin of notum. Scutellum with 2 pairs setae, apical pair long (ca. 2 x length of other pair), cruciate. Wing 0.93 mm long; fairly broad, width less than half the length. Vein C extended to apex of M 1, though obscured because tip of wing is folded; Sc present but very incomplete and faint; R 1 virtually straight; R 2 + 3 parallels R 1, curved costad only at very apex; fork of R 4 and R 5 widely divergent, nearly at right angle; crossveins bm-cu slightly oblique to r-m (not parallel) and proximal to r-m by distance equal to length of r-m; cell dm and crossvein dm-cu present, cell dm large; vein M 2 complete, closing cell dm with crossvein dm-cu; CuA 1 complete; CuA 2 present, complete, forming cell cu p, with incomplete ���spur��� of veins A 1 [+CuA 2?] originating from cell cu p. Anal lobe of wing well developed. Legs long, bristly, with row of stiff setae on ventral margin of hind tibia. ABDOMEN: Very folded and not visible in dorsal view, unclear in lateral view. Type. Sex? (tip of abdomen not preserved), Holotype: SPAIN: ��LAVA, near Pe��acerrada, Escucha Formation, Lower Cretaceous; MCNA 8882. Etymology. From the Latin, concilio, unite separate parts into a whole, connect; in reference to the plesiomorphic features of this genus that link Chimeromyia with other basal Eremoneura. Comments. See discussion of phylogenetic relationships, below., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 50-52, DOI: 10.5281/zenodo.187264

Chimeromyia reducta Grimaldi & Cumming 1999 C. reducta Grimaldi & Cumming 1999: 74. Diagnosis. Immediately distinguished from all other species in the genus by the terminal arista, short posterior ocellar setae; long R 2 + 3 vein gradually sloping towards C; base of M and vein M 2 lost. In Lebanese amber.

Chimeromyia alava Arillo & Grimaldi, sp. n. (Figs. 11, 12) Diagnosis. Similar to C. pilitibia in having crossvein bm-cu slightly proximal to r-m and the male terminalia with a lobed surstylus, but differs by having the surstylus shorter and being trilobed instead of bilobed, R 2 + 3 is shorter and curved more abruptly costad, and the male hind tibia without a brush of long, fine setae. Description. Body length 1.0– 1.45 mm. HEAD: Eyes large, bare, facets not differentiated. Two pairs long ocellar setae present; ocelli on small raised tubercle. Arista situated dorsally on flagellomere one; arista with three articles, basal two very small. Vibrissae present. THORAX: Bristly, with notopleural and supra-alar setae present. Acrostichals in short, single row; one pair of dorsocentals. Scutellum with 2 pairs setae, apical pair longer (this pair cruciate in MCNA 8883). Legs bristly, but without brushes on tibiae or tarsi. Wing: Sc vestigial; C extended to slightly past apex of vein M 1; R 1 meeting C at slightly less than length half length of wing; R 2 + 3 strongly curved toward and meeting C, distances between apices of R 1 and R 2 + 3 only ca. 1 / 2 length of vein R 1 from where it forks with R 2 + 3; vein R 4 + 5 forked with R 4 and R 5 widely divergent, at nearly a right angle; crossveins bm-cu slightly proximal to r-m, separated by distance ca. 0.3 x length of r-m; M 1 straight, nearly parallel to R 5; base of vein M 2 absent, M 2 not reaching wing margin; CuA 1 straight, complete apically. Anal lobe reduced. ABDOMEN: Male terminalia: Fused cerci with pointed apex; epandrium with row of stiff setae on dorsoapical margin; surstylus with three lobes (two lateral ones, one longer, thin mesal one), setulae at apex of only lateral lobes; phallus apically scoop-shaped, slightly S-shaped; postgonites with apex having minute denticles on ventral surface; hypandrium obscured. Female genitalia (MCNA 9318) with cercus large and ovoid in lateral view. Types. All Specimens from SPAIN: ÁLAVA: Peñacerrada, Escucha Formation, Early Cretaceous. Holotype, male, MCNA 9238 (body length 1.45 mm, wing 1.05 mm), with genitalia well preserved. Paratypes: female, MCNA 9318, genitalia well preserved but head and thorax badly preserved (body length 1.25 mm, wing 1.1 mm); male/female, MCNA 8883, a very small specimen (1.00 mm body length), but with venation identical to others in type series; MCNA 8743.2 female, the venation identical to other specimens of C. alava; female, MCNA 8886, head is very distorted and gives an appearance of the arista not being dorsal, venation also identical to others in type series (body length 1.45 mm). All deposited in Álava Museum of Natural Sciences, Vitoria-Gasteiz, Spain. Etymology. Taken directly from Álava, Basque Country of Spain.

Chimeromyia acuta Grimaldi & Cumming 1999 C. acuta Grimaldi & Cumming 1999: 74. Diagnosis. Base of M complete; vein M 2 incomplete, anal lobe and alula lost; readily distinguished on basis of relatively large flagellomere 1, which has an acute ventral margin. Arista dorsal. Sursylus simple (not multilobed), digitiform; postgonite long, apex slightly hooked; apical margin of hypandrium with pair of long thin lobes. In Lebanese amber., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 39-40, DOI: 10.5281/zenodo.187264, {"references":["Grimaldi, D. & Cumming, J. (1999) Brachyceran Diptera in Cretaceous ambers and Mesozoic diversification of the Eremoneura. Bulletin of the American Museum of Natural History, 239, 1 - 124."]}

Chimeromyia intriguea Grimaldi & Cumming 1999 (Fig. 4) C. intriguea Grimaldi & Cumming 1999: 71. Diagnosis (revised). Arista dorsal. Base of vein M (proximal to crossveins r-m and bm-cu) complete, but very faint; crossvein r-m either connected to M or with a slight break near connection; vein M 2 incomplete at proximal end (either evanescent or abruptly), distal end either incomplete or connected to wing margin; CuA 1 complete apically, sometimes faint; anal lobe and alula lost; small spur of vein (CuA 2 + CuP) at base of wing; wing very broad, width approximately 0.5 x the length. Female terminalia simple, with small pair of cerci broadly connected to abdomen. In Lebanese amber. Comments. The diagnosis is slightly revised based on a new AMNH female specimen (Fig. 4).

A new family of eremoneuran Brachycera, the Chimeromyiidae, is proposed for two genera and eight species of a distinctive, monophyletic group of flies in 125–100 myo amber. The new family is related to the Empidoidea and basal Cyclorrhapha. Four new species of Chimeromyia are described: C. pilitibia Grimaldi and Cumming (in Lebanese amber), C. mediobscura Grimaldi and Cumming, C. alava Arillo and Grimaldi (in Spanish amber), and C. burmitica Grimaldi and Cumming (in Burmese amber). A new genus, Chimeromyina Arillo and Grimaldi is also described, for a primitive new species C. concilia (in Spanish amber). New details of these flies are described, particularly of male and female terminalia, and the relationships between this and other eremoneuran families are discussed.

Chimeromyia burmitica Grimaldi & Cumming, sp. n. (Figs. 1���3) Diagnosis. Immediately distinguished from all other species in the family by the loss of vein R 2 + 3; vein M very faint, anal lobe of wing highly reduced, CuA 2 + CuP veins highly reduced to small spur at base of wing; male terminalia with glabrous, sclerotized epandrium; surstyli simple, base enlarged, apical half slender and digitiform. In Burmese amber. Description. A minute fly, body length 1.1 mm. Body apparently yellowish, no apparent color patterns. HEAD: Eyes reddish, without interfacetal setulae, occupying most of lateral surface of head, no differentiation of facets. Two pairs ocellar setae, posterior pair slightly behind posterior ocelli, anterior ocellars slightly posterior to anterior ocellus; ocellar triangle not particularly tuberculate. No setae on frons. One pair of long inner vertical setae present, plus smaller postoculars. Antenna with pedicel cup-shaped, no visible setae; basal flagellomere setulose, lobe-like, with depression and possibly with inserted condyle on mesal surface; rest of flagellum aristate, having two small basal articles and a long, fine, setulose terminal article. Clypeus large, protrudent, apex with pair of thick, stiff setae. Palpus and labellum small; details not visible. THORAX: Mesoscutum with one row of acrostichal setulae; two long rows dorsocentrals flanking acrostichals (slightly larger than acrostichals); each postpronotal lobe with erect seta. Mesoscutellum with single pair of setae (apicals), slightly erect. Wing: Broadly rounded on apex, base narrow, with anal lobe lost. Sc incomplete, apically evanescent; C ending at apex of R 5; R 1 short, apex reaching to midline of C; R 2 + 3 lost; vein R 4 + 5 forked with R 4 widely divergent from R 5, approximately 80 �� divergent from R 5. Vein M very faint, basally and apically evanescent; M 2 present, barely connected to M 1 in short basal fork; crossveins r-m and bm-cu nearly in line, with gap where M intersects. CuA 1 basally sclerotized, portion apical to bm-cu is faint. CuA 2 + CuP highly reduced to small, spur-like vein at base of wing. Legs: Simple, with no distinctive setation or spines, save for row of ca. 6 longer, thin, stiff setae on ventral surface of fore femur. ABDOMEN: fairly long and narrow, tapered apicad, with eight tergites fully visible. Male terminalia: Epandrium short, seemingly heavily sclerotized (darker than rest of sclerites), glabrous save for row of 4 stiff setulae on posterior surface; posteroventral margin with slight notch. Cerci fused medially, long and narrow in lateral view, extended slightly beyond apex of surstyli. Surstyli apparently articulated with epandrium, with basal half broad and apical half slender and digitiform; apparently bare of setulae. Only apex of phallus visible, extended to three-quarters the length of surstyli. Postgonites present, with small dorsal lobe bearing apical seta, several other stiff setae. Hypandrium setulose, simple and lobe-like. Female terminalia (best seen in AMNH Bu 298, where abdomen is cleared, but spermathecal capsules not visible): Cerci lost or not apparent, sternite VIII subtriangular, minute triangular sclerite on ventral surface of apex; no acanthophorite spines. Types. All types in amber from northern Myanmar, Kachin state, near the village of Tanai (ca. 100 km NW of the Mytchkina), coll. 1999 by Leeward Capital Corp. Holotype, male, AMNH Bu 352; Paratypes, two females, AMNH Bu 298, AMNH Bu 504. The holotype is particularly well preserved, though the wings are folded (reconstructed in Fig. 2 B). Etymology. Taken directly from the former name of the Republic of Myanmar., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 36-39, DOI: 10.5281/zenodo.187264

Chimeromyia mediobscura Grimaldi & Cumming, sp. n. (Figs. 5���8) Diagnosis. Base of M (proximal to crossveins r-m and m-cu) virtually lost, fold-like; R 2 + 3 gradually tapered toward C (not abruptly turned); crossveins r-m and bm-cu contiguous or nearly so, but interrupted where M intersects; anal lobe barely evident, no alula, a small spur of a vein at base of wing (CuA 2 + CuP); two pairs of scutellar setae (posterior pair cruciate); arista dorsal; male: surstylus simple and digitiform, postgonites spatula-shaped, hypandrium cleft with apices of lobes pointed, stVIII with row of large setae on apical margin; female terminalia: simple, tapered to narrow apex, cerci present. In Lebanese amber. Description. A small, bristly species, slightly more robust than other Chimeromyia. HEAD: Wider than long; eyes large, bare, facets not differentiated. Ocellar triangle on small, slightly raised tubercle, with 2 long pairs of ocellar setae; posterior ocellars 0.7 x length of anteriors. Pair of long vertical setae, nearly erect; 4���5 postoccipital setae very close to posterior margin of eye. Antenna with flagellomere I setulose, rounded; arista situated near dorsal margin; 3 aristal articles, basal two aristomeres minute, approximately equal in length. Clypeus protuberant; with pair of facial setae present, projecting anteriad, length 0.5 x that of anterior ocellar setae. Palpus and labellum small; labrum visible in one specimen (Az 285), length c. 3 x the width, with pointed apex. Epipharyngeal teeth, if present, not visible. THORAX: Fairly broad, setose. Scutum with one row 6���8 acrostichal setulae; pair of large dorsocentral setae between supra-alars, ca. 5 small dorsocentrals in each row; two supra-alar setae present, two notopleural setae; one erect postpronotal seta; scutellum with apical pair of long, cruciate setae, subapically with smaller, finer pair, 0.7 x length of apicals. Legs long, femora stout (particularly hind femora). Femora with ventral row of ca. 4���6 stiff, fine setae, lengths about equal to width of femur. Legs largely yellowish, apical two tarsomeres dark brown to blackish, very dark in fore tarsi. Hind tibia of male without plumosity, but with dorsal and ventral row of short, stiff setae (Fig. 5 b). Wing with vein C extended to apex of M 1; Sc incomplete, apex evanescent, crossvein h not present; R 1 meeting C near middle of wing; R 2 + 3 nearly straight, sloped gradually toward C; vein R 4 + 5 forked with R 4 and R 5 widely divergent, length of fork significantly less than base of fork; base of M (proximal to r-m and m-cu) fold-like, virtually absent; M 1 parallel to base of fork R 4 + 5; crossveins r-m and bm-cu in line, sometimes with slight break where they would meet M; vein M 2 thin, almost connected distally to wing margin, proximal end evanescent; CuA 1 tapered, small spur of a vein (CuA 2 + CuP) present at base of vein; although anal lobe present but highly reduced. ABDOMEN: Slender, with male genitalia projected posteriad, not dorsoflexed. Male terminalia: Cerci fused at base, narrow and slightly cleft apically, formed hood-like over phallus and surstyli. Surstyli long, simple lobes with 4���5 small setulae at apex; lengths approximately equal to that of phallus. Phallus narrow, apically scoop-shaped; postgonites laterally flat and spatula-shaped, with stiff apical setae on ventral margin, length slightly less than that of phallus. Hypandrium largely hidden by large sternite VIII; apex of hypandrium cleft, with apex of both lobes pointed. Sternite VII with row of ca. 10 large, stiff setae near apical margin. Female terminalia: simple, tapered to narrow apex with cerci present. Types. All types in Lebanese amber. Holotype, male, Az 649; Paratypes: male, Az 440; male, 274; female, Az 285. All in MHNP. Etymology. In reference to the median vein and its faint structure. Chimeromyia pilitibia Grimaldi & Cumming , sp. n. (Figs. 9, 10) Diagnosis. Antennal arista dorsal, not apical; crossvein bm-cu slightly proximal to r-m, not directly in line; base of vein M sclerotized (not evanescent), bisecting cells br and bm; R 2 + 3 abruptly upturned toward C; two pairs scutellar setae, posterior pair cruciate; hind tibia plumose (probably in males only), with 3���4 longitudinal rows of long, fine setae; male terminalia with bilobed (vs. simple) surstylus. Description: Body length 1.40 mm, wing length 1.0 mm. HEAD: Wider than long; eyes large, bare, facets not differentiated. Ocellar triangle on small, slightly raised tubercle, with 2 long pairs of ocellar setae (setal lengths ca. same lengths as dorsocentral setae). 4���5 fronto-orbital setae very close to margin of eye. Antenna with flagellomere I setulose; arista situated dorsally; 3 aristal articles, basal two minute. Face with clypeus protuberant, pair of large, projecting setae. THORAX: Fairly broad, setose. Mesoscutum with row of acrostichal setulae; large pair of posterior dorsocentral setae, smaller dorsocentrals anteriad, several supra-alar and notopleural setae; scutellum with apical pair of long, cruciate setae, subapically with smaller pair (ca. 0.3 x length of apicals). Legs long and slender; most distinctive feature is plumose hind tibia (probably a male character only), with 4 longitudinal rows of long, fine setae. Wing with vein C extended to apex of M 1; Sc not observable; R 1 meeting C at slightly less than basal third of wing; R 2 + 3 curved upward toward C, distance between its apex and apex of R 1 1 / 2 the length of cell r 1; vein R 4 + 5 forked with R 4 and R 5 widely divergent, length of fork slightly less than base of fork; M 1 parallel to base of fork R 4 + 5; crossvein r-m slightly distal to bm-cu, separated by segment less than 1 / 3 length of either crossvein; vein M 2 very diffuse, fold-like, incomplete at both ends; CuA 1 tapered and incomplete apically; veins CuA 2 + CuP not apparent, although anal lobe well developed. ABDOMEN: Slender, with male genitalia dorsoflexed. Male terminalia: Cerci fused, formed hood-like over phallus and surstyli; surstylus with two long lobes, dorsal one slender and digitiform, both lobes with apical setulae (vental lobe with 2���3 setulae on ventral margin). Phallus apically scoop-shaped; postgonite with apical setae, length approximately equal to that of ventral lobe of surstylus. Hypandrium large, convex. Female terminalia simple, apical segment a long and slender cone; cerci small. Type. Holotype, male, LEBANON: ���Ambre de Hammana/Mdeiru, Aptian inf��rieur, Collection Dany Azar, no. 230 ��� on the label. Specimen is in a tiny (4 x 4 x 1 mm) clear yellow chip of amber, mounted in balsam on a microscope slide. The tiny chip includes the pyritized remains of two other complete Chimeromyia (both females) and the head of a fourth individual. In MNHP. Etymology. From the Greek, pilos (hair), in reference to the hind tibia having a feathering of dense, long, fine setae. Comments. The remains of four of these flies in such a minute piece of amber indicate they were either congregating at a resin flux or were swarming. Swarms in empidoids and other Diptera are usually comprised of males, into which females fly to become mated. The eyes of male Eremoneura that swarm are typically holoptic, but those of Chimeromyiidae are not. This species appears to be one of the most primitive species of Chimeromyia, by plesiomorphically possessing a fairly well developed anal lobe at the base of the wing. In those empidoids that have crossveins bm-cu and r-m nearly contiguous (e.g., some extant tachydromiines), bm-cu is usually distal to r-m, unlike C. pilitibia where bm-cu is slightly proximal to r-m. The primitive new genus Chimeromyina, however, shares the same configuration of crossveins with Chimeromyia., Published as part of Grimaldi, David A., Cumming, Jeffrey M. & Arillo, Antonio, 2009, Chimeromyiidae, a new family of Eremoneuran Diptera from the Cretaceous, pp. 34-54 in Zootaxa 2078 on pages 40-48, DOI: 10.5281/zenodo.187264