IRINA BRAKE

Phylogenetic systematics of the (Diptera, Schizopbora)

Entomologica scandinavica Supplement No. 57 now: Systematics & Evolution J1:ntomologica scandinavica Supplements also supplement to: Insect Systematics & Evolution

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© Entomologica scandinavica Supplement 2000 Entomologica Scandinavica Supplement 57

Phylogenetic systematics of the Milichiidae (Diptera, )

IRINA BRAKE 2 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Contents 1. Introduction ...... 3 1.1. Historical review of research on the Milichiidae ...... 4 1.2. Historical review of the relationship between the Milichiidae, Camidae and ...... 5 1.3. ...... 5 1.4. Geographic distribution ...... 8 2. Materials and methods ...... 8 3. Cladistic analysis ...... 14 3.1. Characters used for cladistic analysis ...... 14 3.2. Cladogram ...... 17 4. Morphology and character transfOlmations ...... 17 4.1. Preliminary remarks ...... 17 4.2. Head ...... 19 4.2.1. Postfrons ...... 19 4.2.2. Prefrons ...... 27 4.2.3. Compound eyes ...... 29 4.2.4. Proboscis ...... 29 4.3. Thorax ...... 34 4.3.1. Prothorax ...... 34 4.3.2. Mesothorax ...... 39 4.3.3. Wings ...... 39 4.3.4. Legs ...... 42 4.4. Abdomen ...... 44 4.4.1. Male preabdomen ...... 44 4.4.2. Male pregenital segments ...... 46 4.4.3. Male genitalia ...... 47 4.4.4. Male reproductive system ...... 61 4.4.5. Female ovipositor ...... 61 4.4.6. Female reproductive system ...... 65 4.5. Immature stages ...... 72 4.5.1. Eggs ...... 72 4.5.2. Larvae and puparia ...... 74 4.6. Stem- pattem of Milichiidae ...... 74 5. Phylogeny of the Milichiidae ...... 75 5.1. Phyllomyzinae ...... 75 5.2. ...... 77 5.3. Milichiinae ...... 78 5.4. Minimum age of the Phyllomyzinae, Madizinae, and Milichiinae ...... 79 5.5. Evolutionary "trends" in the biology of the Milichiidae ...... 80 6. Phylogenetic position of the Milichiidae within the Schizophora ...... 82 6.1. Comparison with traditional classifications ...... 82 7. Acknowledgements ...... 83 8. References ...... 84 9. Appendix. Key to genera ...... 89 10. Appendix. Catalogue of world Milichiidae ...... 92 References ...... 109 Index ...... 113

The illustration on the coverof a Litometopa sp. 1. is made by the author. Phylogenetic systematics of the Milichiidae (Diptera, Schizophora)

II(INA BRAKE

Brake, I.: Phylogenetic systematics of the Milichiidae (Diptera, Schizophora). Ent.scand. Supp!. 57:1-120. Lund, Sweden 2000-12-15. ISSN 0105-3574.

A phylogenetic tree based on adult characters is proposed for the Milichiidae. The computer-based cladistic analysis employed 23 taxa and 53 characters. The monophyly of the Milichiidae and of the subfamily Milichiinae is confirmed. However, the subfamily Madizinae in its fonner sense is paraphyletic and has therefore been restricted to the generaDeslI1ometopa, , Litometopa, andMadiza. These genera are more closely related to the Milichiinae than to the generaAldl'ichiom),za, Costalima, genus nov. 1, Micl'osimlls, Neophyllomyza, Param),ia, Phy/lomyza, Stomosis, and Xenophyllomyza. For these latter genera, the new subfamily Phyllomyzinae is established. The sister-group of the Milichiidae is the Chloropidae. It is suggested that the is the sister-group of the Milichiidae + Chloropidae, based on the elongation and coiling up of the spermathecal ducts. The Acartophthalmidae has been used to represent the outgroup for the computer-based cladistic analysis. The morphology and character transformations within the Milichiidae are discussed and the stem-species pattern of the Milichiidae is compared with the stem­ species pattern of the Chloropidae, Acartophthalmidae, , and the Chloropidae family­ group. The main emphasis is laid on the morphology of the labella, ovipositor, female reproductive system, and male genitalia. A catalogue of world Milichiidae and a key to genera are given.

I. Brake, AG Evolutionsbiologie, Institut fUr Zoo logie, Freie Universitat Berlin, Kiinigin-Luise-Str. 1-3, D-14195 Berlin, Germany. t. Introduction One peculiarity of the Milichiidae is that they are very diverse, and for long this has been a source The Milichiidae (Diptera, Schizophora) are of confusion for dipterists: "The form ofthe adults small, mostly black acalyptrate . The family varies from carnid-like genera such as Desmo­ contains about 240 described species in 18 genera metopa Loew, through agromyzid-like genera such (see the catalogue in the Appendix), and is as A ldrichiomyza Hendel, and chloropid-like f011ns worldwide in distribution. such as Madiza [Fallen], to stout-bodied, tachinid­ The behaviour of several species ofMilichiidae like genera such as Eusiphona Coquillett and is vcry specialised. For example, in some species Bilimek. Many characters that are thc adults are myrmecophilous, whilst in some relatively stable diagnostic features in related othcrs they are kleptoparasitic, feeding on the prey families vary within Milichiidae" (J. F. McAlpine or spiders or predaceous . The observation 1989). In spite of this diversity, however, the of' such behaviour raised the question of whether Milichiidae are generally thought to be mono­ ll1yrmecophily and had evolved phyletic (Griffiths 1972, Wheeler 1994). once or several times within the family. To answer The Milichiidae are divided into two subfamilies, this question, a phylogenetic tree of Milichiidae Milichiinae and Madizinae. The Milichiinae are was needed, and so I have studied the phylogeny of thought to be monophyletic, because there are the Milichiidae and have reconstructed a phylo­ several apomorphies for the subfamily. By contrast, genetic tree. I studied over one hundred species in the Madizinae are characterised only by plesio­ total, of which 23 were used in the computer-based morphic characters and have therefore been thought cladistic analysis (see list of species in section 2). to be paraphyletic (Hennig 1971 b, 1973; Wheeler 4 Irilla Brake ENT. SCAND. SUPPL. 57 (2000)

1994). Both the monophyly ofthe Milichiinae and third species, securicornis. Fallen had the paraphyly of the former Madizinae are con­ earlier established the genera Madiza and filmed in the present study. A new monophyletic Phyllomyza in 1810, but without species descrip­ subfamily, the Phyllomyzinae, is established to tions. He placed Madiza in the Oscinides (the include all the genera which have to be excluded modem Chloropidae) andPhyllomyza in Ochtidiae from the Madizinae. (the modem ). The next species The purpose of my studies was to reconstruct a were described by Meigen in 1830. One of these phylogenetic tree of the Milichiidae based on the species, which he placed in the family , is morphological characters of the adults. For the speciosa, and from this the family-name reconstruction of phylogenetic trees, suitable Milichiidae is derived. The other species were representatives ofthe outgroup are essential. In the described in the genus Agromyza, the modern case ofthe Milichiidae, the representative could be . a species from the family Chloropidae because, Schiner (1864) was the first to describe, within according to Griffiths (1972) and Wheeler (1994), the family Muscidae, the "Abtheilung Milichiinae" the Chloropidae are the sister-group of the Mili­ which contained the genera Lobioptera chiidae. However, the stem-species pattem of the (= Milichia), Milichia, and Cacoxenus (now placed Chloropidae is characterised mainly by the reduce in the ). Schiner assignedPhyllomyza tion ofthe characters present in the stem-species of and several "Agromyza" species in his "Abtheilung the Schizophora (Nartshuk 1984) and is therefore Agromyzinae" . not suitable for the determination of character Hendel (1903) was the first to recognise the polarities within the Milichiidae. Since the Milichii­ taxon Milichiidae almost in its modern sense. He dae, Chloropidae, Acartophthalmidae, and Cami­ distinguished two subgroups, which represent the dae have been united by Griffiths (1972) in the subfamilies Milichiinae and Madizinae, but he Chloropidae family-group, I will compare the cha­ used the name "Milichiinae" for the whole group racter states present in the Milichiidae with the and did not name the subgroups. Hendel also added states present in all three of these other families. to the Madizinae the genus Rondani, The Acartophthalmidae resemble the stem-species 1856, which Rondani had placed in his Oscinia pattem of the Schizophora more than the other (the modern Chloropidae) within the Agromyzidae. families ofthe Chloropidae family-group (Griffiths Meoneura is now placed in the Camidae. Appar­ 1972), and so I have used the Acartophthalmidae ently the first to use the name "Madizinae" were as representative of the outgroup in the computer­ Czemy & Strobl (1909), though they also included based cladistic analysis. the genus Rhicnoessa which is now assigned to the In addition to the phylogenetic discussion, the Tethinidae. Duda (1935a) divided the Madizinae evolutionary "trends" in the biology of the into two groups, Madiza-group and Phyllomyza­ Milichiidae will be outlined and the minimum age group, based on the number of orbital setae. These ofthe main groups will be established with the aid generic groups largely correspond to my new of amber inclusions. concept ofthe Madizinae and Phyllomyzinae. How­ ever, Duda's generic groups were never adopted 1.1. Historical review of research on the by subsequent authors. Milichiidae The first extensive study ofthe Milichiinae was by Becker (1907). This paper includes several keys In 1820, the first two milichiid species, Madiza and descriptions of genera and species. The largest glabra and M sordida (= sordida) and most important paper on the Milichiidae was were described by Fallen. In 1823 he described the by Hennig (1937). This comprises keys and ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 5

descriptions of the Palaearctic Milichiidae and a Milichiidae are sister-groups or whether the discussion of the development, geographic distri­ Call1idae are more closely related to one part of the bution and phylogeny of the family. Milichiidae. Based on the shape of the prothorax, Speight (1969) came to the conclusion that the '02. Historical review of the relationship family Milichiidae, which included the subfamily between the Milichiidae~ Carnidae and Call1inae, is probably polyphyletic. In 1971, Hennig Chloropidae accepted Sturtevant's views, placed the Milichiidae and Chloropidae in his Chloropoidea, and treated Thc question of whether the Call1idae is a the Call1idae as a separate family with unknown slIbfamily of the Milichiidae or a separate family sister-group within Chloropoidea (Hennig 1971 a, has for long been a subject for discussion. The 1972, 1973). Since then, the separation of the reason for this is the similarity between the call1id Milichiidae and Camidae, as well as the close genera Meoneura and , and several relationship of the Milichiidae and Chloropidae, Madii'.inac. Both groups are characterised by the has not been questioned. However, the exact rela­ presence of two medioclinate "frontal" setae (see tionship between the Milichiidae, Chloropidae, secli()1l 4.2.1) and two lateroclinate orbital setae. and Call1idae, and the position of these families This condition is unique in the Schizophora. within the Schizophora, has remained a matter for As mentioned above, Hendel (1903) added discussion (Griffiths 1972, J. F. McAlpine 1989). Meli/li'III'1I to the Milichiidae. In 1921 Frey sepa­ ralcd hOlh taxa on the basis of the shape of the 1.3. Biology prohoscis and established the new family Carnidae ror Ihe genera Meoneura and . He placed The biology of the Milichiidae is rather diverse. Ihe Mil ichiidae together with the Chloropidae, Larvae are generally saprophagous and live in ('ollopidac, , and in his decaying plants [Desmometopa (Sabrosky 1983), ( 'ollopiformes, and the Carnidae (which included Enigmilichia (Deeming 1981), Milichia (Malloch Ihe sllhf~lInily Odiniinae) in his Sciomyzaefonnes. 1913, Bohart & Gressit 1951, Wilton 1961), Mili­ S tthseq lIently Hendel (1922-1923) placed the Mili­ chiella (Wilton 1961, Hardy & Delfinado 1980, \·hiidac together with the Agromyzidae, Carnidae Rohacek 1995)]. Some larvae are coprophagous IIIHI as the Milichioidea, next to the and can be found for example in dung in pastures ( 'hloropoidea (which only contained the Chloro­ [Desmometopa singaporensis (Sabrosky 1983), pidae) in his Drosophilomorphae. By contrast, Leptometopa niveipennis (Papp 1992), Madiza SllIrlevant (1925-1926) placed the Milichiidae glabra (Coffey 1966, Papp 1992), :tlld ('hloropidae together in one group, the Chloro­ lacteipennis (Bezzi 1928, Bohart & Gressit 1951)J, pi formcs, after a study of the internal female latrines [Desmom etopa varipalpis (Sabrosky 1983), reproductive system, because "the rudimentary Leptol1letopa coquilletti (Howard 1900, Hennig seminal receptacles with long fine ducts, and the 1956), Milichia pubescens (Deeming & Baez pocket-like ventral receptacle indicate that these 1985)], bat guano [ (Haenni Iwo groups are close to each other." StUlievant did 1988), Pholeomyia dampji (Sabrosky 1959)], and flol examine the Call1idae. chicken manure [Desmometopa spp. (Sabrosky lIennig (1937) added the Call1idae and certain 1983), Madiza glabra (Coffey 1966)]. Larvae of olher genera such as Horaisl11optera (now in the Leptometopa latipes can be found in bird nests Telhinidae) and Risa (now in the ) to (Hicks 1962). Several Desmometopa species are Ihe Milichiidae. In several papers (Hennig 1937, necrophagous and have been found in rotting snails I ()5X) he discussed whether the Carnidae and or on mammal carcasses (Sabrosky 1983). Milichia 6 Irina Brake ENT. SCAND. SUPPL. 57 (2000) piscivorum was reared from decaying fish (Malloch (Sabrosky 1959). Other females were even ob­ 1926). served hopping on to leaves carried by Atta texana Some Milichiidae are myrmecophilous. Larvae foragers and to ride on these leaves into the nest of Phyllomyza have been reared from nests of (Waller 1980). Lasius fuliginosus and Formica rufa, where they One very interesting feature of Milichiidae live in the chambers and galleries at the bottom of behaviour is kleptoparasitism. Desmometopa, Lep­ the nest (Donisthorpe 1927). Microsimus luteus is tometopa, Milichiella, Neophyllol11yza, Paral11yia, known from Crematogaster stoW nests (Aldrich Phyllomyza, and Stomosis species suck at the prey 1926) and Costalima myrmicola from Azteca nests of spiders or predatory insects such as , (Sabrosky 1953). Pholeomyia larvae live in the , Mantidae, or Odonata. In the case of nests ofleaf-cutting ants (Atta texana), where they Desmometopa m-lligrul11 and D. sordida, the prey feed on the detritus ofthe gardens (Sabrosky is always the honey bee Apis mellifera (Mik 1898, 1959, Moser & Neff 1971, Waller 1980). Other Biro 1899, Knab 1915, Kramer 1917, Rabaud Pholeomyia and Eusiphona species have been 1924, Bristowe 1931, Richards 1953, Sabrosky reared from the nests of leaf-cutting bees (Mega­ 1983, Landau & Gaylor 1987). In observations on chile spp.) (Sabrosky 1955, 1982; Teson & other undetermined Desmometopa species, bees Dagobert 1979; De Santis 1981), while Madiza and cicadas were the prey, and females were also glabra was once found in the nest of a solitary attracted to the spray of a Coreidae (McMillan ground-nesting wasp (probably Diodontus tristis 1976). Species of other genera (Leptometopa, Lind.) which stores aphids (de Meijere 1946). Milichiella, Neophyllomyza, ) are mostly Adults ofseveral genera (Desmometopa, Lepto­ attracted to predators feeding on stink bugs metopa, Madiza, Milichia, Milichiella, Pholeo­ (Pentatomidae) or squash bugs (Coreidae) (Frost myia) feed on the nectar of small white or yellow 1913, Robinson & Robinson 1977, Sivinski & flowers, for example Heracleum sphondylium, Stowe 1980, Landau & Gaylor 1987), as well as to Ochradenus baccatus, Solidago canadensis, and the spray of these bugs (Eisner et al. 1991, Aldrich Tanacetum vulgare. In some species adults feed on & Banos 1995). In almost all cases it is only the the honeydew of aphids or other Homoptera females that are kleptoparasitic. In some cases a [Desmometopa (Herzig 193 8),Milichia (de Meijere close association between milichiid and predator 1909, see below, Deeming & Baez 1985), has been postulated, because it was observed that Milichiella (1. C. Deeming, pers. comm.)] or are the "rides" on the predator for some time, fed by ants: the adults of Milichia dectes, M. staying with the one predator rather than changing proectes, and M. prosaetes are known to solicit between different predators (Biro 1899, Robinson regurgitated food from Crematogaster ants (Collin & Robinson 1977). Milichiidae are not attracted to 1921, Farquharson 1921). M. brevirostris licks dead bees or bugs as such, but at least the dead anal secretions from the ant Dolichoderus predator has to be there as well (Biro 1899). The bituberculatus or collects honeydew directly from distribution of Milichiidae is distinctly clumped. the Homoptera (Membracidae) which are attended Consequently they are seldom collected, but then by the Dolichoderus ants (de Meijere 1909). in large numbers. For this reason Sivinski & Stowe As yet no feeding association between Pholeo­ (1980) have suggeste'd that either some spiders (or myia species with myrmecophilous larvae and ants predaceous insects) or their prey are more attractive has been observed, but these flies use the ants than others or the clumping results "from a instead as a guide to theirnests: Pholeomyia females simultaneous emerging brood that locates a host were observed following leaf-cutting ants (Trachy­ near the site of ecclosure." It was observed on myrmex) , probably to find the nest entrance several occasions that the flies "clean" the mouth- ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 7

I=l I=l 0 0 I=l I=l '5b '5b 0 0

Genus nov. 1 xx

Tab, I, Geographic distribution ofMilichiidae, x - one species, xx - more than one species. (The total number of described sJlecics is added in brackets behind the genus name), parts of the predator (McMillan 1976, Sternberg or meadows, in wadis, at the edges of forests, 19l)1), and this could be the reason why these inside forests, in the forest canopy, in stables or predators do not fend off the flies. houses, or even in caves. However, they do not The habitats of Milichiidae are diverse. Adults seem to be attracted to coastal habitats or to other call be collected in open landscapes, such as steppes places near water. 8 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

1.4. Geographic distribution USNM National Museum of Natural History, Washington, USA The geographic distribution of the Milichiidae ZMAN Zoologisch Museum Amsterdam, can be seen in Tab. 1 (and is listed for each species Netherlands in the catalogue in the Appendix, p. 91). In general, Milichiidae prefer high temperatures and the Most of the material studied was preserved in greatest number of species is therefore found in 70% ethanol (all material from UBI, part of the tropical and subtropical regions. This is especially material from NMW.Z, and part of my own true for the Milichiinae: in Central and Northern collection); the rest was pinned. The material from Europe there is only one species, Milichia ludens. the University of Bielefeld and my own specimens were determined by me, whilst the other material 2. Materials and methods has been identified by various dipterists. Some specimens were dissected (see list of species). The About 90 species of Milichiidae (see list of dissected parts were macerated in heated 90% species) and a few Chloropidae, Acartophthalmidae lactic acid and stored in euparal. and Carnidae for outgroup comparison were studied For scanning electron microscopy studies, with a Leitz stereo microscope, and with compound alcohol-fixed specimens were transferred in 10% microscopes (Leitz Diaplan and Zeiss Axioplan). steps to 96% ethanol and critical point dried with This material was borrowed from the following carbon dioxide (CPD 030, Balzers Union). Dry persons or institutions: specimens were sputtered with gold (SCD 040, Balzers Union) and studied with a Philips SEM ANIC Australian National Insect Collection, 515. Canberra For histology, dissections were embedded in BRAKE Private collection of!. Brake, Gelmany resin (Kulzer Histo-Technique) and sectioned 2-3 DEI Deutsches Entomologisches Institut, [lm thick with a R. Jung microtome. The sections Eberswalde, Germany were stained with Toluidine blue. ETH Eidgenossische Technische Hoch- The cladistic analysis was carried out using the schule Zilrich, Switzerland computer software Hennig86 (Farris 1988), in HOFFEINS Private collection of C. & H. W. conjunction with DADA (Nixon 1995) and Hoffeins, Germany CLADOS (Nixon 1994). HU Museum filr Naturkunde, Berlin, I have studied one species of each genus in Germany detail and have used its characters for the computer- ISMAY Private collectionofJ. W. Ismay, Great based cladistic analysis. Where possible, I have Britain studied at least two further species, to ensure that NMW.Z National Museum and Galleries of the characters selected were not autapomorphies at Wales, Cardiff, Great Britain the species level but were common characters for OZEROV Private collection of A. L. Ozerov, species-groups or for the whole genus. When I use Russia generic names in the following text, I will only POINAR Private collection of G. Poinar, Cali- refer to the species studied by me for a certain fornia character (see list of species) or to data available SMN Staatliches Museum fur Naturkunde from the literature. Stuttgart, Germany In the case of two genera, Milichia and Pholeo- TAU Tel Aviv University, Israel myia, I have included two species from each genus UBI University of Bielefeld, Germany in the character matrix. I have includedPholeomyia ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 9

1 Siphonellopsinae Oscinellinae Chloropinae

median vertical seta longer than lateral vertical seta costa reaching R4+5 only anteroventral corner of epandrium projecting 2 posterior notopleural setae

basi phallus large femoral organ 2 humeral setae katepisternal setae lost

orbital setae numerous and short

precoxal bridge

only one costal break

crossvein bm-cu lost anal vein and anal cell lost IChloropidae I tibial organ three pseudotracheae short, slender duct between spermathecal duct and capsule

2 Neomeoneurites Hemeromyia Meoneura Camus

T610st crossvein bm-cu lost apical half of M 1 very weak R4+5 and M1 diverging abdominal pleural membrane with fine setulae, each with a sclerotised base two lateroclinate upper and two medioclinate lower orbital setae

female cerci fused I Carnidae I bases of antennae recessed into deep fossae one additional medioclinate orbital seta below the three lateroreclinate setae

Figs. 1-2. - 1. Phylogenetic tree of the Chloropidae (modified after Andersson 1977 and Nartshuk 1984 and 1987). - 2. Phylogenetic tree of the Carnidae (modified after Hennig 1972 and Grimaldi 1997). Black hatchmarks indicate apomorphies; characters added by me are marked with dark shading. nigricosta (Hendel, 1932) in the character matrix knowledge of the family, I have assumed that all because the generic position of this species has genera of the Milichiidae are monophyletic, but I been disputed (see section 5.3). The reason for have not checked the monophyly of each genus. including two Milichia species in the character This was beyond the scope of the present work. matrix is that the genus is probably para- or poly- Genera whose monophyly is doubtful (for example phyletic. In general, and based on my previous Milichia and Leptometopa) are discussed indivi- 10 Irina Brake ENT. SCAND. SUPPL. 57 (2000) dually. tatives ofthe outgroups used in the cladistic analysis. In sections 4.2-4.4, which deal with the My discussion of the stem-species pattern of the morphology ofthe Milichiidae, I am only describing Chloropidae is based on the papers of Andersson structures which are meaningful for the phylogeny (1977) and Nartshuk (1987). In the case of the ofMilichiidae or for the position ofthe Milichiidae Acartophthalmidae, I assume thatAcartophthalmus within the Schizophora. I make no claim to be nigrinus represents the stem-species pattern of the exhaustive. Furthermore, certain data for some family because the ACaliophthalmidae comprises taxa are missing, because I have not dissected only one genus, , with about 4 specimens of species where only few specimens species. In addition, I discuss the stem-species exist, e.g. Enigmilichia dimorphica, Microsimus pattern of the Carnidae and of the Chloropidae luteus, and Pholeomyia nigricosta. Costalima is family-group, because the 4 families Milichiidae, the only genus that I have not studied at all. Chloropidae, Acartophthalmidae and Carnidae Prosaetoll1ilichia (de Meijere, 1909) is not were united in the Chloropidae family-group by discussed because I recently synonymised it with Griffiths (1972). The Carnidae have not been Milichia (Brake 1999). included in the cladistic analysis because support In sections 4.2-4.4, I not only describe the for the inclusion ofthe Carnidae in the Chloropidae character states present in the Milichiidae but I also family-group is weak (see section 6). A phylo­ describe the ground pattern of the stem-species genetic tree for the Chloropidae and Carnidae can (stem-species pattem) of the Chloropidae and the be found in Figs. 1 and 2. Acartophthalmidae, because these are the represen-

List of species studied

The following list contains all the species of any dissections. Milichiidae, Chloropidae, Acartophthalmidae, and Abbreviations: 1M - imago (male), IF - imago Carnidae that I have studied. Species used for the (female), P - proboscis, M- male genitalia, MR­ computer-based cladistic analysis are underlined. male reproductive system, 0 - ovipositor, FR - The total number of described species in each female reproductive system, L-Iegs, D -digestive genus can be found in Tab. 1. Except where system, Pu - puparium, E - eggs. otherwise stated ("IF only" or "1M only"), I have Dissections: Italics: sections, bold: SEM studied the imago of male and female in addition to dissections.

Species: Locality, Collection: Dissections:

Milichiidae:

Aldrichiomyza A. agromyzina (Hendel, 1911) North America, USNM P,M A. elephas (Hendel, 1913) Taiwan, USNM A. longirostris HendeL 1931 Israel, TAU P, M, 0, FR, 1M

Desmometopa D. ciliata Hendel, 1919 Western Australia, UBI (X649) IF D. inaurata Lamb, 1914 Western Australia, UBI (X632) ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 11

D. interji-ontalis Sabrosky, 1965 Cape Verde Is., UBI (X380) D. m-nigrum (Zetterstedt, 1848) Canary Is., UBI (X386) Greece, UBI (5.4. 85) °1M, IF, 0 Israel, BRAKE M,FR,D D. saguaro Sabrosky, 1983 Arizona, BRAKE D. sordida (Fallen, 1820) Germany, UBI (L283-3l2) P,P,M Germany, BRAKE MR, FR, 1M, M, IF, 0 D. varipalpis Malloch, 1927 Gelmany, BRAKE 1M, IF, Pu D. cf. meridionalis Argentina, UBI (X 1066) P,O

Enigmiliclzia E. dimor12.hica Deeming 1981 Nigeria, NMW.Z IF only ?E. sp. 1 Uganda, UBI (X1259-Ra35) IF only

Eusiphona E. mira Coquillett, 1897 Texas, HU IF only lLm-J. Western Australia, ANIC P,M,O

Leptometopa L. latipes (Meigen, 1830) Germany, BRAKE FR, 1M, IF France, UBI (X814) P,M,O L. mallochi Sabrosky, 1989 Australia, ANIC M L. nivei12.ennis (Strobl, 1900) Gelmany, BRAKE P,M,P,IM L. ru:ftji-olls Becker, 1903 Israel, BRAKE O,FR Greece, UBI (XnO) P L. sp. 1 Israel, BRAKE FR L. sp. 2 Uganda, UBI (X1259-Ca14) ° Litometopa L. glabrifi~ons Sabrosky, 1965 Tanzania, SMN (holotype) L. ?glabrifi'ons -~ ~ Uganda, UBI (X1259-TnI2, P, M, O,FR - Ra3) IM,IF L. sp. 1 Tanzania, TAU P,M,F L. sp. 2 Uganda, TAU P,M

Madiza M. glabra Fallen, 1820 Gelmany, BRAKE P, P, MR, M, M, FR, D, 1M, IF, MR, E M. pachymera Becker, 1908 Israel, TAU

Microsimus M. luteus Aldrich, 1926 Bolivia, USNM (paratype) IF only 12 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Milichia M argyratoides Collin, 1921 Uganda, UBI (X1259-Ca8) M brevirostris (de Meijere, 1909) Java, ZMAN (lectotype) IF only M distinctipennis Deeming, 1981 Ivory Coast, UBI (X1320) P,M,O,FR M Jarquharsoni Collin, 1921 Ivory Coast, UBI (X1320-38) FR M Jormicophila Deeming, 1976 Uganda, UBI (X1259-Ra14) P M myrmecophila (de Meijere, 1909) Java, ZMAN (lectotype) P, 1M, IF, M M ludens (Wahlberg, 1847) Norway, BRAKE Switzerland, ETH M° M. speciosa Meigen, 1830 Romania, DEI P, M, O,FR M. sp. 1 Uganda, UBI (X1259-Ca18) IF only M. sp. 2 Uganda, UBI (X1259-Ca13) IF only M. sp. 3 Spain, UBI (X1205-11) IF only, FR

Milichiella M. lacteipennis (Loew, 1866) Israel, BRAKE MR, M, FR, 1M, IF,E M.sp;l Turkey, UBI (X477) M M. sp. 2 Venezuela, UBI (X1285) M,FR M. sp. 3 Cuba, UBI (X780) M. sp.4 Venezuela, UBI (X1293) ° Neophyllomyza N. acyglossa (Villeneuve, 1920) Germany, UBI (L1550-14, L 1553-12) P, 0, FR, P Switzerland, ETH P,M N. sp. 1 Uganda, UBI (XI259-Tn12) P,M,O,IM N. sp. 2 Borneo, NMW.Z IM,IF N. sp. 3 Japan, UBI ° Paramyia P. inconspicua de Meijere, 1916 Rwanda, UBI (X1006-Cg5,6,15) P. nitens (Loew, 1869) Brazil, UBI (X493) ~ Uganda, UBI (X1259-Tn14) P, M, 0, FR, 1M, IF P. spp. 2-7 Borneo, NMW.Z P,P,M

Pholeomyia P. longiseta (Becker, 1907) Argentina, UBI (X1077) M P. nigricosta (Hendel, 1932) Bolivia, SMN, (holotype) IF only ~ Jamaica, UBI (L1890) P, M, 0, FR, P, 1M P. sp. 2 Brazil, UBI (X493) P. sp. 3 Venezuela, UBI (X890) ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 13

Phyllomyza P. beckeri Kramer, 1920 Great Britain, BRAKE P. equitans Hendel, 1919 Germany, UBI (L1552) P. jlavitarsis (Meigen, 1830) Gelmany, UBI (L2241) P. securicornis Fallen, 1823 Gelmany, UBI (L1914-16, P,P,P L283-312, K895) M,O,FR,IM P. sp. 1 Borneo, NMW.Z P,P,L, P. sp. 2 Uganda, UBI (XI259-Ra3) M P. sp. 3 Uganda, UBI (XI259-RaI3) o

Stomosis S. innominata (Williston, 1886) Panama, USNM S. vittata Malloch, 1925 Western Australia, UBI (X663) P,P,O,M,IM ~ Western Australia, UBI (X649) P, M, FR, P, 1M, M S. spp. 2-10 Australia, ANIC P,M,O

Ulia U. poecilogastra Becker, 1907 Pern, HU (holotypeJ IF only Costa Rica, DEI P,M,FR

Xenophyllomyza X deserticola Ozerov, 1992 Turkmenistan, OZEROV P, M, FR, IF ?X. sp. 1 Western Australia, UBI (X669) P, P, M, IF

Genus nov. 1 sp. 1 I)()i \{l Borneo, NMW.Z P,O,FR :jli \ Genus nov. 1 sp. 2 U>"".(]", v·,." Borneo, NMW.Z P,O

Amber inclusions:

?Milichia cf. ludens Dominican amber, POINAR IF only Milichiella sp. 5-7 Dominican amber, POINAR 1M, IF Phyllomyza ?hurdi Dominican amber, POINAR 1M, IF Phyllomyza sp. 4-5 Dominican amber, POINAR 1M only Phyllomyza sp. 6 Baltic amber, HOFFEINS 1M only ?Pseudodesmometopa sp. I Baltic amber, HOFFEINS IF only

Chloropidae:

Apotropina (Siphonellopsinae) A. senilis (Duda, 1930) Venezuela, UBI (XI279) FR A. ? jilscipleuris (Becker, 1911) Papua New Guinea, ISMAY P,M,O,FR A. longepilosa (Strobl, 1893) Portugal, ISMAY P, M, 0, FR, 1M 14 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Oscinellinae Germany, Israel, BRAKE P,M,FR Anatriehus sp. 1 Ivory Coast, BRAKE P

Chloropinae Germany, Israel, BRAKE P,MR,FR

Acartophthalmidae:

Aeartophthalmus A. bieolor Oldenberg, 1910 Gelmany, HU P,FR A. nigrinus (Zetterstedt, 1848) Sweden, UBI P,M,FR,O

Carnidae: Hemeromyia H. sp. 1 Israel, BRAKE P, MR, M, FR, IF Meoneura M. lamellata Collin, 1930 Germany, BRAKE P,M, FR, Pu M. ji-eta Collin, 1937 Germany, BRAKE M M. laeteipennis (Fallen, 1823) Germany, BRAKE P,M,MR,MR, FR, FR, 1M, IF M. vagans (Fallen, 1923) Germany, BRAKE MR,FR M. flavifacies Collin, 1930 Germany, BRAKE MR,FR,

Neomeoneurites N. ehilensis Hennig, 1972 Chile, SMN (holotype)

3. Cladistic analysis Head Cl Upper orbital seta latero-reclinate (0); latero­ The data set for the computer-based cladistic clinate (1); medio-reclinate (2); absent (3). analysis comprised 53 characters and 23 taxa, C2 Middle orbital seta latero-reclinate (0); including the Acartophthalmidae as representative lateroclinate (I); latero-proclinate (2); absent of the outgroup. The Chloropidae were treated as (3). an ingroup. All characters were treated as unor­ C3 Lower orbital seta latero-reclinate (0); latero­ dered. The cladistic analysis was carried out using clinate (1); latero-proclinate (2); absent (3). the exact algorithm option "ie" ofHennig86, which C4 Number of frontal setae: zero (0); two (1); is guaranteed to find the most parsimonious three (2). cladograms. C5 Postocellar setae diverging (0); strongly converging or cruciate (1); parallel or slightly 3.1. Characters used for cladistic analysis converging (2). C6 Proclinate setula between supra-antennal seta The precise character distribution can be seen in and eye margin absent (0); present (1). the data matrix (Tab. 2). C7 Lunule bare (0); with one pair of setae (1); ~ ;l (/Jn Character number 5 10 15 20 25 30 35 40 45 50 ~ Genus nov, 1 sp, 1 o 0 0 000 1 0 ? 1 1 000 1 0 1 0 0 0 0 0 0 0 0 0 200 005 1 001 000 0 0 1 0 1 0 0 ? t! (/J Costafima myrmicola 3 ? 1 0 0 0 0 ? 0 ? ? ? ? 0 ? ? 0 0 0 0 0 0 ? 0 ? ? ? 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? sa Microsimus lute us 0 0 0 0 0 1 2 ? ? ? 0 0 ? 0 0 0 0 0 0 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? >-0 Phyllomyza securicomis 1 0 0 0 0 1 2 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 0 0 0 0 0 0 0 0 0 0 0 !:""' '-" Neophyllomyza acyg/ossa 3 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 3 2 2 0 2 0 0 0 0 0 ---l ~ tv Paramyia sp, 1 2 1 1 0 0 0 0 0 0 0 0 3 0 0 0 0 2 1 0 0 0 0 0 0 0 0 0 0 3 0 2 0 2 0 0 0 0 0 o o o Stomosis sp, 1 1 1 0 0 0 0 0 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 0 3 0 1 0 1 0 0 0 0 0 '---' A/drichiomyza longirostris 3 0 2 0 0 0 0 0 0 0 1 0 3 0 0 0 0 2 0 0 0 0 0 0 0 0 1 0 3 2 0 0 0 0 0 0 0 0 Xenophyllomyza deserticola 3 0 2 1 0 0 0 0 0 0 1 0 3 0 0 0 1 0 1 2 1 0 - 0 0 0 0 0 1 0 3 2 1 1 0 0 0 0 0 Leptometopa niveipennis 2 2 0 0 0 0 0 0 0 0 0 0 2 0 0 3 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 2 0 0 0 0 0 0 0 0 0 Madizaglabra 202 2 000000000020000000-00000001220001000000 Desmometopa sordida 2 1 2 0 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 4 2 Lilometopa ?glabrifrons 3 0 3 1 2 0 0 0 a a a 0 a a 3 a a a a 0 a 0 0 a a a a 0 a 0 a 0 2 2 a 0 a 4 2 1 1 Enigmilichia dimorphica 3 a 2 2 0 a O? a a 0 2 0 a 0 0 0 0 0 0 0 0 0 ? ? 0 0 ? ? ? ? ? O? 1 0 0 0 ? 0 ? Eusiphona sp, 1 - 2 0 a 0 0 0 a 2 0 0 0 0 0 0 0 a 0 0 2 1 0 2 0 a 3 0 0 0 0 0 Milichia distinctipennis 3 a 2 2 1 2 0 1 0 a 0 0 a 2 0 0 1 0 0 a 0 0 a 0 a 0 1 2 a 0 0 ? 0 ? a 0 0 2 0 2 2 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 0 0 2 2 0 0 0 0 3 0 0 0 0 0 Ulia poecilogastra 303 1 200 001 100 1 0 1 0 ? 0 0 1 101 0 100 0 1 1 003 1 2 1 1 1 ? ? ? ? 1 0 ? 0 0 a >-0 Milichia speciosa 2 a 2 020 0 a 1 a ? 0 a 0 ? 2 001 1 000 1 000 1 1 000 1 2 1 4 1 000 1 100 ,E[ Pholeomyia sp, 1 20221 0 0 101 0 0 0 202 000 000 1 000 2 0 0 004 2 1 0 0' \fQ Pho/eomyia nigricosta 2 022 2 0 0 0 ? 2 1 0 ? ? ? 0 ? 2 0 1 201 000 0 0 0 ? ? ? ? ? ? ? ? ? ? a 0 0 ? ? ? ? ? ? ? (1) ~ Stem species pattern Chloropidae o 2 2 0 a 0 0 0 0 0 0 002 000 300 3 0 0 0 000 001 000 a 0 0 0 0 0 1 0 0 000 1 0 0 0 0 0 o >-+, Acartophthalmus nigrinus o 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 000 0 2 0 0 0 a 0 a a a 0 0 0 0 0 0 2 0 002 1 2 1 0 0 a 0 0 0 ~ 0' s:p; p:J (1)

Tab, 2, Data matrix for cladistic analysis, (- = non-applicable character; ? = missing data,) For explanation of characters, see text '-" 16 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

with two pairs of setae (2). (0); bare (1); with three or four strong setae C8 Vibrissal angle present and vibrissa below (2); covered with hairs (3). lower margin of eye (0); vibrissal angle C24 Number of katepistema I setae: one (0); more obsolescent and vibrissa above lower margin than one (1). of eye (1). C25 Subcostal break in costa present (0); develo- C9 First flagellomere in males as large as in ped into a notch (1); absent (2). females (0); first flagellomere in males larger C26 Costa reaching Ml (0); reaching R4 + 5 only than in females (1). (1). CI0 Eye about as high as wide or up to 1.5x as C27 R4 + 5 and Ml parallel (0); converging (1); high as wide (0); more than 1.5x as high as diverging (2). wide (1). C28 Crossvein dm-cu present (0); absent (1). Cll Frons in males as wide as in females (0); C29 Distal margin of anal cell (CuA2) rounded narrower than in females (1). (0); CuA2 meets anal vein in an acute angle C12 Posterior eye margin without notch or (1 ). emargination (0); with notch (1); with C30 Shape of wing oval (0); triangular (1). emargination (2). C31 Hind femur without strong seta (0); with Cl3 Maxilla with ventral appendage and lacinia strong seta anteroventrally (1). present (0); ventral appendage absent and C32 Sensory area on hind tibia absent (0); present lacinia strongly reduced in length or absent (1). (1). C33 First tarsomere of hind leg ventrally without C14 Palpi short and clavate (0); long and thin (1); a row of setae (0); with a posterior comb of long and thick (2). setae (1); with several rows of setae (2). CIS Outer surface of labella glabrous (0); pube- scent (1). Male abdomen C16 Number of pseudotracheae: more than four C34 T2-5 not silvery microtomentose (0); comp- (0); four (1); three (2); two (3). letely or partly silvery microtomentose (1). CI7 Medial pseudotracheae separate (0); con- C35 Tergites slightly bent atthe sides (0); strongly verging at tip (1). bent ventrally at the sides and sometimes CI8 Labella, when open, together up to twice as with a crease (1). long as broad (0); more than 2x as long as C36 S5 as long as or longer than S4 (0); S5 much broad (1). shorter than S4 or absent (1). Cl9 Pseudotracheal ring tips at distal end ofmedial C37 Surstylus and epandrium not fused (0); fused pseudo trachea in females as long as the other (1 ). ring tips (0); longer than the other ring tips C38 Surstylus with narrow tip (0); rounded and with narrow tip (1). apically (1); spoon-shaped (2); with two lappets (3); with appendage (4); notched (5). Thorax C39 Postgonite present (0); absent (1). C20 Basistemum small (0); reduced to a narrow C40 Phallapodemic sclerite not fused with triangle (1); large and V -shaped (2); with hypandrium (0); fused with median part of pre coxal bridge (3). hypandrium (1); fused with bases of C21 Proepimeral seta absent (0); present (1). pregonites and hypandrium (2). cn Number of postpro notal setae: one (0); more C41 Median part of hypandrial arc present (0); than one (1). absent (1). C23 Anepistemum with hairs and up to two setae C42 Distiphallus long, flexible, coiled, and ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 17

pubescent (0); short and glabrous (1); form­ medio-reclinate, or absent ing a slender tube, which is widened in the *C2-0/1 Middle orbital seta latero-reclinate or middle (2); sock-shaped (3); fODning a slen­ lateroclinate der tube bearing fine hairs (4). *C7 -0/1 Lunule bare or with one pair of setae C43 Distiphallus without sclerotisations (0); with *C 15-0/1 Outer surface oflabella pubescent or posterior sclerotised stripe at base (1); with a glabrous stripe on each side at base (2). *C23-1/3 Anepis~ernum bare or covered with hairs Female abdomen *C40-0/2 Phallapodemic sclerite not fused with C44 T7 complete (0); divided longitudinally (1); hypandrium or fused with bases of divided halfway longitudinally (2). pregonites and hypandrium C45 T8 complete (0); divided longitudinally (1); *C47-0/1/4 S8 complete, longitudinally divided, divided halfway longitudinally (2). or fonning two strips, each with a row C46 S7 complete (0); divided longitudinally (1); of setulae divided halfway longitudinally (2). *C48-1/2 Ring of barbed spines between seg­ C47 S8 complete (0); divided longitudinally (1); ments 7 and 8 absent or between S7 divided halfway longitudinally (2); absent and S8 (3); forming two strips, each with a row of *C49-0/1 Secondary ovipositor absent or present setulae (4). *C50-0/1 Ovipositor long or shorter than width C48 Ring of barbed spines between segments 7 of abdomen and 8 present (0); absent (1); present between *C51-01l Supra-anal plate with one pair of S7 and S8 (2). setulae or bare C49 Secondary ovipositor absent (0); present (1). *C52-0/1 Cerci long or short C50 Ovipositor long (0); shorter than width of The reasons for these alterations will be abdomen (1). discussed in the following sections. In most cases C51 Supra-anal plate with one pair of setulae (0); I have suggested that certain character states bare (1). evolved earlier in the phylogeny ofthe Milichiidae C52 Cerci long (0); short (1). because, in my opinion, it is more probable that C53 Spermathecal duct coil tightly wound (0); these characters evolved once and were reduced loosely wound (1). several times, than that they evolved several times independently. In consequence, the number of 3.2. Cladogram steps needed for the tree increases. This is also the reason why Hennig86 does not produce this tree. The cladistic analysis with Hennig86 using the exhaustive search Hie" found 13 most parsimonious trees of158 steps (consistency index: 0.56; retention 4. Morphology and character trans­ index: 0.73). Only in fourofthese trees were all the formations branches supported, and so I discarded the other 4.1. Preliminary remarks trees. Upon closer inspection I removed some poorly supported branches in the four trees, and the 1) In sections 4.2--4.4 I am combining descrip­ resulting polytomous trees were identical (fig. 3). tions ofthe morphology and ofthe character trans­ In addition, I altered the point of origin of character fODnations within the Milichiidae, although the states in a few cases: polarity of character transformation results from *Cl-l/2/3 Upper orbital seta lateroclinate, the cladistic analysis and not from the morphology 18 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Phyllomyzinae Madizinae Milichiinae I I Jl? co 0 C/) C/) (.) .~ .~ C/) ~ 'e ~C/) t::: Q) co .§ E: Q t::: "b .~ .~ ~ C/) Q) C/) Jl? 0 Q) .g t::: t::: .Ql 0 ~ .8- t::: t::: 0 .g (.) "b OJ 'E Q) (.) t::: .S? C/) t::: 0 Q) ~ ci.. ~ ~ co co C/) .8- C/) co ';:: C/) C/) (.) co Q .~ :§- (1j C/) .Ql ~ E .2l Q) t::: co (.) .2l .$2 ci.. C/) ~ co Q ci.. (.) OJ (j) t::: .§ ~ ~ co ~ (j) (.) ci.. ci.. E: ..Q :§ Q) (J) co E' C/) co C/) (j) ~ Q .8 ~ ~ :> E: Q Q) co (.) .~ .~ ell 0 co ~ 0 co E: .8 ~ t::: (1j -0 :S ~ .~ ~ Q) OJ :S ::::: CD ~ ..c: (J) c 's., C/) .$2 E: 0 E: E: '0.. .S 'S, ..c: 0 0 0 ell (j) ~ C/) E: ..c: 0 ..c: E: co ..c: .~ ·92 .~ 0 0 g. :l Q E: g. (.) ..c: ..c: Q.. ..c: "- ~ ~ Q E: .8- (.) ~ ~ 1:: c C/) e E: .8 :g C/) C/) (.) .S? 0 .Q c (.) ~ 0 ~ t::: Q Jg 0 co "- (J) 0 (J) .8 :§ Q) Q) ~ ..c: ..c (.) ell (9 () ~ it <:: ~ C/) ~ <::( -J ~ Q lLJ ~ ~ ::> ~ it Q 0 <::( 0 45-1 1-2 6-0 -0 1~3 7~(j 6,Q, H~26~1 28-'1 23-3 3-2 3,3: 22-1 9-1 .27~2 44-1 38-1 32-1 6'0' 37_1.28-1 45-1 47~0 38-1 ,. J2:J 'r~ii 1:3: 10- 16_0 24-1 43-2 46-1 .48,1 47-1 ~7-g ~!J,O 48:0 2-1 38-1 46.,1: 12-1 24-1 ltM, 14-1 50-0 i49,Q 24-147:1 :49-0 15" 43-1 37-1, ~C:j, 51-0 50-0 25-0 ;48,1 50-0 i8~1 47-1 46-1, 2-0, 52-0 51 -0 38-149'051_0 24-1 .4~c1· 44-1 3-2, 52-0 19-1 ,44,2 51-0 52-0 34-1i411~0 47-3 52-0 43- 42-4 2-3 14-2 38-4 ,H), 53-1 24-1, 48;9 4!F1 eS,cj',27-2, All,.D 49'0 as-o, 43-2 50-0 38-2 '51-0 51-0 52-0 20-1, 52-? 45-2, 47-2

43-1 42-2

2-0,3-2,10-1,27-1

8-1,11-1,29-1

21-1,33-UI8:1. 17-1

1-1,2-1,3-1,4-1,6-1,7-1,16-1 '39-1

Fig. 3. Phylogenetic tree of the Milichiidae. Shaded hatchmarks indicate character changes that either reverse or evolved' at least twice on the tree; black hatchmarks indicate changes that do not reverse and evolved only once; white hatchmarks indicate plesiomorphic character states; the character numbers of reductions are marked with dark shading. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 19 itself. This is done to give a better understanding of with the fact that the head has to be elastic when the the transformations. To follow the phylogenetic ptilinum is evetied (Hennig 1973). The only parts explanations more easily, the cladogram in Fig. 3 which are strongly sclerotised are the ocellar should be consulted. triangle and the lateral orbital plates, which are 2) In addition to the characters which were used situated next to the compound eyes (PI. lA). In J. in the cladistic analysis, I am also describing several F. McAlpine (1981), an orbital plate is called other characters. These characters were not used in "fronto-orbital plate" if it extends to the anterior the cladistic analysis either because they would margin of the frons, and "orbital plate" if it is occur only once in the matrix or because they are restricted to the upper half of the frons. However, difficult to define on account oftransitional stages. these plates are homologous structures and so I am 3) I am using the symbol" + "to indicate sister­ using only the name "orbital plate". group relationships. For example A ldrichiomyza + According to Hennig (1958), the orbital plate Xenophyllomyza stands for the monophyletic group convergently became smaller along different consisting of the genera Aldrichiomyza andXeno­ evolutionary lines of the Schizophora, so that the phyllomyza. Brackets indicate the relationships orbital plate with the orbital setae is restricted to within the monophyletic group. For example, Sto­ the upper half of the postfrons. In the lower half of mosis+ (Aldrichiomyza+ Xenophyllomyza) means the postfrons, the frontal vitta extends from eye that these three genera constitute a monophyletic margin to eye margin (holometopy), and only a group in which Stomosis is sister-group of Aldri­ nalTOW strip along the lower eye margin remains chiomyza + Xenophyllomyza. more strongly sclerotised. This strip may widen 4) For the morphology I am following the secondarily and the lateral row of frontal hairs may terminology ofJ. F. McAlpine (1981) except where develop into strong medioclinate setae, which are stated otherwise. For the chaetotaxy, only long called frontal setae and look exactly like orbital setae are counted and not short ones. setae. This secondarily widened plate is called the 5) Hypotheses for the function of certain frontal plate. This situation (schizometopy) evolved characters are typed in Petit, because I have not independently in the , , and studied the function of any characters or the Milichiidae, and can be recognised by a clear behaviour of the flies. separation between the orbital and frontal plates in some species (PI. lA). It must be differentiated 4.2. Head from the ancestral shape, in which the orbital plate reaches the anterior margin ofthe frons. In some of The main parts ofthe head capsule that I studied these ancestral shapes (present in the Agromyzidae, are the frons anteriorly, compound eyes and genae Camidae, Odiniidae), the anterior orbital setae are laterally, and proboscis ventrally. The frons is medioclinate (PI. 4A) and are called frontal setae divided into a dorsal postfrons ("frons") and a by some authors, though they are true orbital setae. ventral prefrons ("face") by the insetiion of the Further characteristic setae are located on the antennae. The different inclinations of the head ocellar triangle ( ocellar setae) and on the vertex setae are shown schematically in Fig. 4. (medial and lateral vertical setae). The vertical setae are traditionally called "inner and outer" 4.2.1. Postfi-ons vertical setae, but I prefer to use the terms "medial and lateral" as proposed by Freidberg & Beschovski The middle of the postfrons in the Schizophora (1996). Behind the ocellar triangle are converging is more or less membraneous and is called the or diverging postocellar setae, and on the frontal frontal vitta. A membraneous frons may cOlTelate vitta are interfrontal setulae, hairs, or both. 20 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A B

if s --~=I'---":--' frs=-----t-"~

D

I

poe s E \

orbs~ __,,---.

PI. 1. A-B. Postfrons, frontal view (setae shown on the left half, cuticula structures on the right half). - A. Pilyllol11yza securicornis, male. - B. Stomosis villata, male. - C. Genus nov. I sp. I, male, head, frontal view. - D. Stomosis sp. I, male, head, lateral view. - E. Milichia myrmecophila, male, head, frontal view (after Brake 1999). - F. Elisipilon(J mira, female, head, lateral view (after Sabrosky 1987a). Abbreviations: ar- arista; fr pl- frontal plate; fr s - frontal seta; fr vit­ frontal vitta; if s - interfrontal setnla; I v s -lateral vertical seta; lun -lunule; m v s - medial vertical seta; oc s - ocellar seta; oc tr - ocellar triangle; orb pl- orbital plate; orb s - orbital seta; pip - palpus; poe s - postocellar seta; prob - proboscis; pst s - peri stomal seta; pv s - para vertical seta; sant s - supra-antenna I seta; sl- setula between supra-antennal seta and eye margin; vi - vibrissa. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 21

Orbital setae (Cl-3) secondarily enlarged so that it is as long as the In the stem-species pattern of the Milichiidae lower orbital setae in Pholeomyia, Ulia sp., and in there are three orbital setae, which are apo­ some Milichiella species (PIs. 2A, 3E). morphically lateroc1inate (Fig. 5). In my opinion, I am homologising the lateroc1inate upper orbital 3 orbital setae which are latero-reclinate belong to seta with themedio-rec1inate seta because in groups the stem-species pattern ofthe Chloropidae family­ with 3 lateroc1inate orbital setae there is no seta or group, because this character state is present in the setula between the uppermost orbital seta and the Acartophthalmidae (PI. 9B) and several other medial veliical seta, but in several genera with 2 schizophoran families. orbital setae (Desmol11etopa, Leptometopa, Madiza, In the stem-species pattern of the Carnidae Paramyia) (PI. 2B-D) there is a small additional

convyrging cruc,iate latero-reclinate / /medio-reclinate ~--'-.l..\---,...... reclinate

Fig. 4. Schematic drawing of the different inclinations of the head setae.

(present inNeomeoneurites andMeoneurites), there medio-reclinate seta above the orbital setae. In are also 3 latero-reclinate orbital setae, but there is addition, there is a large medio-reclinate upper an additional fourth orbital seta below the others orbital seta present in Pholeomyia and Milichiella which is slightly medioclinate (Hennig 1972). In lacteipennis (PIs. 2A, 3E). Hemeromyia, Meoneura, and Camus there are 2 The inclination of the two lower lateroc1inate lateroclinate upper orbital setae and 2 medioclinate orbital setae also changes within the Miliehiidae. lower orbital setae (PI. 4A). In the stem-species pattern the middle one is slightly In the stem-species pattern of the Chloropidae rec1inate and the lower one is slightly proelinate. there are 3 orbital setae, which in the ancestral state However, in the stem-species of A ldrichiomyza + are either all proc1inate (Nartshuk 1987), or the Xenophyllol11yza, inMadiza, and in the Milichiinae upper one is latero-rec1inate and the lower ones the middle orbital seta is mainly reelinate and only latero-proc1inate (Andersson 1977) (PI. 3F). slightly lateroclinate, and the lower orbital seta is The stem-species pattern of the Milichiidae, mainly proelinate and only slightly lateroclinate with 3 lateroc1inate orbital setae, is present in (PI. 2A, D). Where there is only one orbital seta, as Costalima, genus nov. 1,Microsimus,Phyllomyza, in Litometopa and Ulia poecilogastra, this is latero­ and Stomosis (PI. lA-C). Within the Milichiidae, reclinate and is therefore probably homologous the upper orbital seta became smaller and medio­ with the middle orbital seta (PI. 2E). It follows that reclinate convergently in Paramyia and in the the upper and lower orbital setae are lost in stem-species ofthe Milichiinae + Madizinae (stem­ Litometopa and Ulia poecilogastra. species G). The upper orbital seta is lost secondarily In the type specimen of Ulia poecilogastra and in Aldrichiol11yza + Xenophyllomyza, Enigmili­ in the other specimens studied, there is only one chia, Litometopa, Milichia distinctipennis, Neo­ rec1inate orbital seta. According to A. L. Serra phyllomyza, and Ulia poecilogastra (Fig. 5; PIs. (pers. comm.), however, there are 3 setae (2 rec1inate 2E, 4E). In contrast to this, the upper orbital seta is and 1 proc1inate) in the Ulia poecilogastra specimen 22 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A B

if s _-+.f--b:F-\''1'-.

if ~tr-~,LJ-..!,----+

C D

o ______m V>-::ipf/"~Ol orb s \0; \ "I .--- fr s r fr s 1:-- 1 0 \ ! 0 0 I Y1~ I

E

PI. 2. A-D. Postfrons, frontal view (setae shown on the left half, cuticula structures on the right half). - A. Pholeomyia sp. I, male. - B. Des1I1ometopa varipaipis, female. - C. Leptometopa iatipes, female. - D. Madiza giabra, male. - E-F. Litometopa sp. I, male, head. - E. Frontal view. - F. Lateral view. Abbreviations: ar - arista; cpist - epistoma; 1st flm­ first flageliomere; fr s - frontal seta; gen - gena; if s - interfrontal setula; if str - interfrontal stripe; I v s -lateral vertical seta; lun -lunule; m v s - medial vertical seta; oc s - ocellar seta; orb s - orbital seta; ped - pedicel; poc s - postocellar seta; pst s - peri stomal seta; sant s - supra-antennal seta; sl-setula between supra-antennal seta and eye margin; vi - vibrissa; vi ang - vibrissal angle. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 23

Phyllomyzinae Madizinae Milichiinae I I

(1) ~ :> C/) (1) E: 0 co :::J ~ 0 c .S E: ~ If) ~ C/) 0 ..c: :::l ~ Q c C/) e(,) ~ 0 (J.) 0 ..c: Q) (') 0 ~ Q <:

mid orb s absent

D F

two pseudotraeheae c

mid orb s reclinate low orb s proclinate

upp orb s medio-reclinate

4 pseudotraeheae 3 pseudotraeheae I : 2 frontal setae mid orb s proclinate I . all orb s lateroelinate low orb proclinate

poe s diverging

- more than 4 pseudotracheae - no frontal setae ~~. ~~~~~. ~-~} - all orbital setae latero-reclinate

Fig. 5. Phylogenetic tree of the Milichiidae with a depiction of the transformation of six characters of the head (CI-C3 inclination of orbital setae, C4 number offrontal setae, C5 inclination of postocellar setae, C 16 number of pseudo tracheae ). Shaded hatchmarks indicate character changes that either reverse or evolved at least 2x on the tree; black hatchmarks indicate changes that do not reverse and evolved only once; white hatchmarks indicate plesiomorphic character states; reductions are marked with a darker shading. Abbreviations: low -lower, mid - middle, orb - orbital, poc - postocellar, s - setae, upp­ upper. 24 Irina Brake ENT. SCAND. SUPPL. 57 (2000) ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 25 that he studied. Either Sena studied a new species parallel (PIs. 4A, 9B). Since the inclination of the of Ulia or the number of orbital setae is variable in postocellar setae is variable in other Schizophora, Ulia poeeilogastra. In either case there are probably the stem-species pattern ofthe Chloropidae family­ 3 orbital setae in the stem-species pattern of Ulta, group cannot be determined at present. and 2 were lost within the genus. Within the Milichiidae, the stem-species pattern Within the genus Costalima, the middle latero­ is represented in the Phyllomyzinae, in which the clinate orbital seta became smaller. I suggest that postocellar setae are cruciate or so strongly 3 orbital setae are in the stem-species pattern of converging that the tips of the setae meet (PI. lA, Costalima because, according to A. L. Serra (pers. C). A synapomorphy for the Madizinae and comm.), there is an undescribed Costalima species Milichiinae are parallel or only slightly converging with 31ateroclinate orbital setae. In the type species, postocellar setae (PIs. IE, 2B-E), but in some C. myrmieola, however, there are only 2 orbital Pholeomyia species they are strongly converging setae. In my opinion, and on the basis of the secondarily (PI. 2A). position of the setae in a drawing by Serra, the middle orbital seta was reduced in this species. Interfrontal setae (C6) The configuration of orbital and frontal setae in In the stem-species pattern of the Milichiidae, Eusiphona differs from all other Milichiidae: all there is a paired row of interfrontal setulae on the the setae together fonTI a uniform row of 8-10 frontal vitta (PI. 2B). This is a plesiomorphic reclinate setae (PI. IF). This is an apomorphy for character state for the Milichiidae, because in the the genus Eusiphona. stem-species pattern of the Chloropidae there is probably a row of 3-4 interfrontal setulae too Frontal setae (C4) (NatiShuk 1987), and in the Carnidae there is a row In the stem-species pattern of the Milichiidae of 2-3 interfrontal setulae (PI. 4A). The stem­ there are apomorphically 2 medioclinate frontal species pattern of the Acartophthalmidae is un­ setae. These frontal setae are not homologous with known: inAeartophthalmus bieolorthe interfrontal the medioclinate setae on the frons of the carnid setulae are unevenly distributed on the frons (fig. genera Hemeromyia, Meoneura, and Carnus, be­ 189 in Hennig 1958) and inA. nigrinus there are no cause the medioclinate setae in Carnidae are orbital or up to 3 interfrontal setulae present (PI. 9B). setae. Within the Schizophora, interfrontal setulae which Within the Milichiidae, 3 frontal setae are an fonTI 2 rows on the frons are present for example in apomorphy for the stem-species of Pholeomyia and Tethinidae. The rows may (PI. 2A). In the Miliehia speeiosa-group the frontal therefore be plesiomorphic for the Chloropidae setae are lost (PI. IE). family-group. The anteriormost interfrontal setula is called Postocellar setae (C5) the supra-antennal seta. It stands next to the ptilinal In the stem-species pattern of the Milichiidae, fissure, is pro- and medioclinate, and is often the postocellar setae are cruciate, because this is stronger than the other interfrontal setulae. Between the character state present in the Chloropidae (PI. the supra-antennal seta and the eye margin there is 3F). In the Acartophthalmidae, the postocellar a further small proclinate setula at the anterior setae are diverging and in the Carnidae they are margin of the frons in the Milichiidae (PIs. IB, C,

PI. 3. A-F. Head, frontal view. - A. Genus nov. 1 sp. 1, male. - B. Leptollletopo iatipes, female. - C. Desmometopa varipaipis, female. - D. Madiza glabra, male. - E. Milichiella iacteipennis, female. - F. Apotropina longepilosa (Chloropidae), male. Abbreviations: epist - epistoma; fr s - frontal seta; if str - interfrontal stripes; lun -- lunule; orb s­ orbital seta; poe s - postoeellar seta; sl- setula between supra-antennal seta and eye margin. Scales 0.1 mm. 26 Irina Brake ENT. SCAND. SUPPL. 57 (2000) ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 27

2B-E, 3A-E). This setula is an apomorphy for the morphically 2 pairs of setulae on the lunule. Milichiidae because it is absent in the Chloropidae, Acatiophthalmidae,andCamidae. Within the Mili- 4.2.2. PreFons chiidae the setula is secondarily lost in Aldri­ chiomyza, in Microsimus, and in the stem-species The lower margin of the prefrons (epistoma) is ofa group comprising the Milich ia speciosa-group warped forward in Leptometopa, Litometopa, + Pholeomyia and maybe Ulia (Pis. IE, 2A). Madiza, Paramyia, andXenophyllomyza (Pis. 2E, In Desmometopa and some Pholeomyia and F, 4C). In Leptometopa the epistoma is triangular Phyllomyza species, the row of interfrontal setulae and nearly meets the long lunule dorsally (PI. 3B). is emphasised by sclerotisations at the base of the Vibrissae (C8) setulae and the development of microtrichiae. In In the stem-species pattem of the Chloropidae such cases the rows are called interfrontal stripes family-group, the vibrissa lies below the lower eye (PI. 2B). Interfrontal stripes are an apomorphic margin or level with it. This character state is character for Desmometopa, but according to present in the stem-species pattern of the Sabrosky (1983) there are several unnamed Milichiidae too (PI. 4C, G, H). Neotropical species without interfrontal stripes In the stem-species of the Milichiinae, the area (but with interfrontal hairs) that may belong to of the vibrissal angle is smaller, and the position of Desmometopa. According to Sabrosky, this con­ the vibrissa has shifted to a position above the dition may be a reduction ofthe interfrontal stripes lower eye margin (PI. IF). This condition has been (see section 5.2). caused by the enlargement ofthe eyes in the stem­ Lunule (C7) species ofthe Milichiinae (see section 4.2.3). The lunule is well developed in most Milichiidae and is quite large inLeptometopa and some Milichia Antennae (C9) species (Pis. IE, 3B). However, in several species In the stem-species pattem of the Milichiidae of the Phyllomyzinae the lunule is partly covered and of the Chloropidae family-group, the first by the anterior margin of the frons. flagellomere is relatively small and rounded in The lunule is bare in the stem-species pattem of both sexes. However, in males ofCosta lima, Micro­ the Chloropidae family-group and in most Schizo­ simus, Neophyllomyza, Paramyia spp.,Phyllomyza, phora. In cases where setae are present on the and Stomosis sp. (PI. 4F) the first flagellomere is lunule in other schizophoran families, there are, so larger than in females and often elongate. The far as I know, several setae. The existence of enlargement of the first flagellomere may have exactly one pair of setae on the lunule is therefore evolved once in the stem-species of the Phyllo­ an apomorphic character of the Milichiidae (Pis. myzinae, and was secondarily lost in genus nov. 1, lA, B,E; 2A-D; 3E). Within the Milichiidae, these several Paramyia species, most Stomosis species, setae are lost in genus nov. 1, Aldrichiomyza + and Aldrichiomyza + Xenophyllomyza. On the Xenophyllomyza, Neophyllomyza, Pholeomyia other hand it is possible that the enlargement nigricosta, Ulia, and most Paramyia species (PI. evolved in Costalima + (Microsimus + Phyllomyza) 4E). In Milichia distinctipennis there are apo- (stem-species D) and convergently in Neo-

PI. 4. A.lvfeonellra lacteipenllis (Carnidae), female, head, frontal view. - B-D. Head, lateral view. - B. Aldrichiol11yza longirostris, male. - C. Xenophyllol11yza deserticola, female. - D. ?Xenophyllomyza sp. 1, female. - E. Aldrichiol11),za longirostris, male, head, dorsal view. - F. Phyllol11yza secllricomis, male, right antenna, lateral view of medial side. - o­ R. Head, lateral view. - O. Leptol11etopa latipes, male. - H. Desmol11etopa varipalpis, male. Abbreviations: epist - epistoma; 1st flm - first flagellomere; fr s - frontal seta; orb s - orbital seta; pIp - palpus; poe s - postocellar seta; prob - proboscis, vi - vibrissa. Scales 0.1 mm. 28 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

lab

E "" ... ~; -Iac-&

PI. 5. A. Phoieomyia sp. 1, female, proboscis, lateral view. - B. Meoneura iacteipenl1is (Carnidae), female, proboscis, lateral view.- C-F. Maxilla (withoutpalpus). - C. Phyl/omyzasecul'icornis, dorsal view. - D-F, lateral view.- D.Anatrichus sp. 1 (Chloropidae). - E. Acartophthaimus nigrinus (Ac3Itophthalmidae). - F. Meaneura iacteipenl1is (Carnidae). - O­ R. Desmometopa sordida, female. - O. Labella, terminal view. - H. Section (positioned at interrupted line in 0). Abbreviations: fulc - fulcrum; hyphar - hypopharynx; lab -labium; lac - lacinia; lbl- labella; lbr - labrum; pip - palpus; prem - prementum; pstr - pseudotrachea; pstr rt - pseudotracheal ring tips; spi - spine; stip - stipes; v app - ventral appendage. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 29 phyllomyza and withinParamyia andStomosis. In hovering or swarming males is known for some my opinion, the latter hypothesis is more probable groups of , many Orthorrhapha, Platy­ because it is more parsimonious. pezidae, , , Syrphidae, and for Calyptratae (Downes 1969). Even in swarming 4.2.3. Compound eyes (CIO-12) species of Ephemeroptera, , Lepido­ ptera, and Trichoptera the compound eyes are fre­ In the stem-species pattern of the Milichiidae quently larger in males than in females (J. F. McAlpine and of the Chloropidae family-group, the eyes are & Munroe 1968). more or less circular in lateral view (PIs. ID; 2F; 4 B-D, G, H). In the stem-species ofthe Milichiinae, A diagnostic character for Milichiella and Ulia the eyes are apomorphically enlarged in width and is a notch in the middle ofthe posterior eye margin especially in height, and are thereby more than (Fig. 6A). Moreover, there arePholeomyia species l.5x as high as wide (PIs. IE, F, 3E). Only in Ulia with a slight notch or an emargination in the lower must this enlargement have been secondarily lost. half of the posterior eye margin, for example Pholeomyia nigricosta (Fig. 6B). This emargination was the reason why Becker (1907) separated the B genusPseudomilichia fromPholeomyia. However, there are transitional stages between notch, emar­ gination, and a straight eye margin. This character alone is not useful for the separation of genera, and so Hendel (1932) synonymised Pselldomilichia with Pholeomyia (see section 5.3). Fig. 6. Schematic drawing of the lateral view ofa compound eye. - A. Posterior eye margin with notch. - B. Posterior eye margin with cmargination. 4.2.4. Proboscis

Correlated with the enlargement of the eyes, the The mouthparts of most higher Diptera consist genae are often nanow so that the eyes take up of a tube-like sucking apparatus, the proboscis nearly the whole lateral side ofthe head. In addition (PIs. SA, B; 7A). The main parts of the proboscis there is a sexual dimorphism, because the eyes of are the basal fu1cmm, and the labium with the the males are even larger than the eyes of the distal labella. The proboscis is connected to the females. This further enlargement results in the head capsule by the clypeal membrane. The fulcmm postfrons being much narrower in males than in is surrounded by this membrane. Anterolaterally at females, which is another apomorphy for the the distal end of the fulcmm are the maxillae. Milichiinae. Cardo, stipes, and lacinia are fused to each other, but the cardo and stipes are enclosed within the The enlargement of the eyes and narrowing of the clypeal membrane, while the lacinia is external. frons probably is connectcd to the swarming The stipes bears the maxillary pal pus at its proximal behaviour in males, because the condition, in which tip and in most acalyptrate flies there is a ventral the upper part ofthe eyes nearly meet in the mid-line, appendage at the middle ofthe stipes (Frey 1921). seems to be an adaptation to the recognition and The labium consists ofthe proximal prementum capture of females in flight (Downes 1969). It is and the distal labella. The prementum forms a known from Milichia jlll11icostata and several trough on the ventral side, in which the hypopharynx Milichiella species that the males swarm in sunlight and labmm are situated. At its distal end it bears the (see section 4.4.1). The enlargement of the eyes in paired labella. The inner surface of the labella is 30 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

o E

pstr -=--f'ffil-+'l::jj I

PI. 6. A-F. Labella, ten11inal view. - A. Phyl/omyza securicornis, male. - B. Genus nov. I sp. 1, female. - C. Milichia distinctipennis, female.- D. Stomosis vitlata, female. - E. Neophyllomyza acyglossa, fe;11ale. - F. Leptometopa rujiji-ons, male. - G-H. Section through labella. - G. Stolllosis vitfata. - H. Neophyllomyza acyglossa. Abbreviations: pstr - pseudo trachea; pstr rt - pseudo tracheal ring tips. Scales 0.1 111111. furnished with small sc1erotised grooves, remi­ tracheae. The pseudotracheae radiate from the niscent of tracheae, which are called pseudo- terminus ofthe food canal and are used to distribute ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 31 saliva and to suck up liquids. They are braced by a whereas in the Chloropidae there are species in series ofsclerotised rings, the tips of which can be which the lacinia is very Sh011 and species in which of different shapes. Zaitsev (1982) distinguished it is completely absent. two basic types of pseudotracheal ring tips: the The reduction ofthe lacinia occurs convergently dentate type (rounded or flattened tips) and the in several schizophoran families, for example in spinose type (elongated and pointed tips). the , Ephydridae, , Platy­ Intermediate shapes of the ring tips are possible stomatidae and . Both lacinia and ventral (Elzinga & Broce 1986). appendage are absent in the and Calyp­ Because the proboscis is Sh011 and not geniculate tratae (Frey 1921). The presence or absence of the in most Schizophora and in the Acartophthalmidae, ventral appendage has not yet been used in I think that this is also the shape of the proboscis in phylogenetic systematics and the character is poorly the stem-species pattel11 ofthe Chloropidae family­ known in most families. Consequently it is not group. The proboscis of the Call1idae is apomor­ known whether the ventral appendage evolved phi cally highly specialised (PI. 5B): the prementum once or several times in the Schizophora. If it is strongly sclerotised and bulbous, and the labella evolved once, then this would suppOt1 the hypo­ are very Sh011 and inconspicuous. thesis that the loss of the ventral appendage is a An apomorphy for the Milichiidae + Chloropidae synapomorphy for the Milichiidae and Chloropidae. is a slightly elongated and geniculate proboscis, If the ventral appendage evolved several times which means that the labella are folded back. I within the Schizophora, thcn it could be possible have not included this character in the cladistic that the ventral appendage is absent in the stem­ analysis, because it is very difficult to decide species pattern of the Chloropidae family-group whether a proboscis is slightly elongated or not and evolved convergently in the ACat10phthalmidae elongated, and if it is only slightly elongated, and within the Call1idae. whether it is geniculate or not. The geniculation is The palpi are Sh011 and clavate in the stem­ only easy to see in long probosces (Pis. ID, F; 4C, species pattern of the Chloropidae family-group. D). In species with a strongly elongated proboscis, The plesiomorphic state of the palpi is represented the labium with the labella and sometimes the in the stem-species pattern of the Milichiidae. fulcrum is elongated, but never the stipcs. Within the Milichiidae, in the stem-species of Aldrichiomyza + Xenophyllomyza the palpi are Maxillae (C13-14) apomorphically long and thin (PI. 4C, D) and in In my opinion, a visible lacinia and a ventral Phyllomyza + Microsimus they are long and thick. appendage are part of the stem-species pattel11 of the Chloropidae family-group, because these Labella (ClS-19) characters are present in the Acat10phthalmidae In most Milichiidae the outer surface of the and in the stem-species pattern of the Call1idae (PI. labella is glabrous and there are only a few setulae 5E, F). The maxilla lacks lacinia and ventral (PI. 7 A). In my opinion, glabrous labella are a appendage only in the carnid Neomeoneurites plesiomorphy and part ofthe stem-species pattel11 (Hennig 1972), which can be interpreted as an of the Milichiidae, because the outer surface of the apomorphy. labella is also glabrous in the Chloropidae. A synapomorphy for the Milichiidae and The stem-species pattel11 for this character in Chloropidae is the absence ofthe ventral appendage, the Chloropidae family-group is uncertain: the which was for the first time studied by Frey (1921) labella in the Acartophthalmidae are covered with (PI. 5C, D). In addition, the lacinia is very small or short hairs on the outer surface, and so this may be absent: in the Milichiidae it is always absent, the stem-species pattel11 ofthe Chloropidae family- 32 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

PI. 7. A. Phyllomyza securicornis, female, proboscis, lateral view. - B-C. Labella, terminal view; B. Phylloll1yza securicornis. - C. Pholeomyia sp. 1, male. - D. ?Xel1ophyllomyza sp. 1, tip oflabellum, dorsolateral view. - E-F. Labella, terminal view. - E. Desmometopa ciliata, female. - F. ?Xel1ophyllomyza sp. I. - G. Stomosis sp. 1, tip oflabella, terminal view. - H. Desmometopa ciliata, female, tip of medial pseudotrachea, lateral view. Abbrev,iations: Ibl-Iabella; Ibr-Iabrum; pip - palpus; pstr - pseudotrachea; pstr rt - pseudotracheal ring tips; spi - spine; 0 Ibl- outer surface of labella; wt - wart­ like structure. Scales 0.1 mm.

group and glabrous labella an apomorphy for the labella are differently structured, and it has not Milichiidae + Chloropidae. However, this character been investigated in other Schizophora. is not applicable to the Carnidae, in which the Within the Milichiidae, the outer surface of the ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 33 labella is covered with fine ShOli hairs in all the tracheae are the 2 medial ones. Furthermore, the 2 Milichiinae that I studied, except for Ulia. However, medial pseudotracheae in genus nov. 1 sp. 1 are the proboscis of Enigmiliehia was not studied. (So much stronger in build than the outer ones (PI. 6B). this character may either be an apomorphy for the As a model, this situation could indicate an Milichiinae or may have evolved after Enigmiliehia evolutionary transition between species with 4 and branched off.) 2 pseudotracheae, though the species itself does The number ofpseudotracheae on each labellum not represent the transitional stage (genus nov. sp. in the stem-species pattem of the Milichiidae is 2 has equally strong pseudotracheae, and both four, because this character state is present in the species are closely related to Phyllomyza). Milichiinae, in the Madizinae, and in part of the The reduction from 4 to 2 pseudo tracheae is Phyllomyzinae. In the Chloropidae there are three probably an adaptation to the elongation of the pseudotracheae (Frey 1921, Graham-Smith 1930, proboscis. On the other hand, it is possible to Elzinga & Broce 1986). In the Acartophtha1midae, elongate the labella without reducing the number there are 12 pseudotracheae in Aeartophthalmus of pseudotracheae, as can be seen in Leptometopa nigrinus and 9 in A. bieolor. In the Camidae there rufifrons (PI. 6F). This species has a very long are 5 pseudotracheae in Neomeoneurites (Hennig proboscis with 4 pseudotracheae. 1972), 4 in Hemeromyia and Meoneura, and 3 In my opinion, the fact that the reduction of the rudimentary pseudotracheae in Cal'l1us (Meoneura number of pseudotracheae is not necessarily and Carnus: Frey 1921). correlated with the elongation of the proboscis In my opinion, there are more than 4 pseudo­ supports the hypothesis that the reduction to 2 tracheae in the stem-species pattem ofthe Chloro­ pseudotracheae evolved only once, in stem-species pidae family-group, because this is the case in E. most Schizophora (Frey 1921) as well as in the In species with an extremely long proboscis (up Acartophthalmidae. Consequently, 4 pseudotra­ to lOx as long as broad) the pseudo tracheae are less cheae are an apomorphy for the Milichiidae, 3 than half as long as the labella. In these cases the pseudotracheae are an apomorphy for the Chloro­ pseudotracheae are situated at the distal end ofthe pidae, and 4 or 5 pseudo tracheae are an apomorphy labella and the food is passed along a common for the Carnidae (Fig. 5). channel (PI. 7F). In the stem-species pattern of the Chloropidae family-group and ofthe Milichiidae (present in the The function of the elongation of the proboscis is Milichiinae and Madizinae), the pseudotracheae unknown. One possibility is, that the length of the terminate separately at the margin of the labella proboscis is cOlTelated to the length of the calyx of (PIs. 5G; 6C; 7C, E). As an apomorphy for the flowers in nectar-feeding flies. I observed Mili­ Phyllomyzinae, the 2 medial pseudotracheae are chiidae with a short proboscis, like DeslIlollletopa convergenting at their tips (PIs. 6A, B, 7B). In spp., Leptometopa spp., Madiza gfabra, and Mili­ stem-species E the labella are apomorphically very chieffa lacteipennis, taking nectar on small yellow long (more than 2x as long as broad) and there are or white flowers. However, there is no reference in only 2 converging pseudo tracheae (PIs. 6D, E; 7D, the literature about milichiids with long proboscis F, G). The question arises as to which of the visiting flowers. Therefore the hypothesis that the original 4 pseudotracheae were lost in the stem­ long proboscis is an adaptation to nectar feeding can species of this group. In my opinion, the outer 2 not be supported at the moment. pseudotracheae were lost, because when only 2 Myrmecophily is known only from species with pseudotracheae are present they are convergenting, shOli proboscis and kleptoparasitism from species and in Phyllomyza the convergenting pseudo- with short or long proboscis. According to Sivinski 34 Irina Brake ENT. SCAND. SUPPL. S7 (2000)

& Stowe (1980) a long proboscis may aid klepto­ Pholeomyia, as well as in the males of some parasitic milichiids on spiders in penetrating silk Desmometopa species (D. ciliata, D. sordida, D. enclosing a spider's prey. Alternatively, a long varipalpis), the ring tips at the tip of the medial proboscis may be advantageous to safely suck up pseudotrachea are longer than the other ring tips fluids near the dangerous mouthparts of a feeding and have a narrow tip (Pis. SG; 7E, H). In the same spider. species the medial pseudotrachea is more strongly The poor knowledge of the biology of Milichiidae sclerotised than the outer three (PI. SH). and the difficulties to observe the behaviour of these The labella of the females of Desmometopa, little flies render it impossible at the moment to Litometopa, and Pholeomyia, and the males of cOlTelate changes in the morphology of mouthparts some Desmometopa species, are similar in several to the biology. characters: (1) the medial pseudotrachea is more strongly sclerotised than the others, and (2) its ring The pseudotracheae in the Milichiidae are 0.01- tips are spinose distally. FurthelIDore, there is in 0.03 mm broad, regardless oftheir length. Only the Desmometopa andPholeomyia (PI. 7H) (3) a spine, weak outer pseudotracheae of genus nov. 1 sp. 1 and (4) a wart-like structure with sensory papillae are slightly narrower. In most Milichiidae the near the tip of this pseudotrachea. In Pholeomyia pseudotracheae are broader basally than distally. sp. 1 there is also a small field of denticles next to InStomosis the surface between the two pseudo­ the spine and the wart-like structure. These tracheae is strongly sclerotised and smooth, and characters are probably used for rasping, and merges into the inner margin ofthe pseudotracheae evolved convergently in Desmometopa + Litome­ (PI. 6G). Consequently there are ring tips only at topa and inPholeomyia. There is another character, the outer margin of the pseudotracheae in this the secondary ovipositor, which is present in genus (PI. 7G). This could be an apomorphy for Desmometopa, Litometopa, and Pholeomyia (see Stomosis or could have evolved within the genus. section 4.4.S). It is possible that the proboscis is In other genera the surface between the pseudo­ used as an aid during oviposition, for example in tracheae is not so strongly sclerotised and both rasping plant tissue. sides ofthe pseudotracheae have ring tips (Pis. SH, 6H,7D). The ring tips of the pseudotracheae differ in 4.3. Thorax shape and size. In the stem-species pattern of the 4.3.1. Prothorax Milichiidae, they are probably round because this is the case in most species of the Phyllomyzinae The dipteran prothorax comprises pronotum, and Milichiinae (PI. 7B). In the Chloropidae, the -pleuron, and -sternum. The setae on the postpro­ pseudotracheal ring tips are either short and pointed notum, which belongs to the pronotum, and the (spinose), or rounded, and in the acartophthalmid setae on the proepimeron, which belongs to the Acartophthalmus nigrinus they are roughly propleuron (PI. 8A), are phylogenetically important rounded. for the Milichiidae. The prosternum includes pre­ Within the Milichiidae, in genus nov. 1 sp. 1, the and basisternum . .I use the terms pre- and basi­ ring tips of the outer pseudotracheae are rounded sternum in the sense of Speight (1969), in contrast and the ring tips of the medial pseudotracheae are to 1. F. McAlpine (1981), who uses the term spinose, with 1-2 tips. In most species of the "prosternum" for the basisternum. The basisternum Madizinae the ring tips are spinose at the base of all lies between the insertion of the fore coxae. The the pseudotracheae. presternum is located in the cervical membrane, In females of Desmometopa, Litometopa, and anterior to the basisternum. It is in contact anteriorly ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 35

Phyllomyzinae Madizinae Milichiinae I I I :::J §

c E

basisternum large and V-shaped

pre coxal bridge I anepisternum with hairs

- basisternum small - anepisternum with hairs and setae ~_~=====~l---_-.J - one katepisternal seta - R4+5 and M1 parallel

Fig. 7. Phylogenetic tree of the Milichiidae with a depiction of the transformation of four characters of the thorax (C20 shape ofbasisternul11, C23 setation of anepisternum, C24 number ofkatepisternai setae, C27 inclination ofR4 + 5 and M i). Shaded hatchmarks indicate character changes that either reverse or evolved at least 2x on the tree; black hatchl11arks indicate changes that do not reverse and evolved only once; white hatchl11arks indicate piesiol11orphic character states; reductions are marked with a darker shading. 36 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A hd B

ex bs sec sci E

..'b~......

ex bs bs G ..... f/!JJ.

bs

PI. 8. A-C. Prothorax, fronto-ventral view. - A. Desmometopa (redrawn from Speight 1969). - B. Madiza (redrawn from Speight 1969). - C. Litometopa ?glabrifi·ol1s. - D-G. Basisternum. - D. Desmometopa saguaro. - E. Phylloll1yza securieornis. - F. Neophyllomyza aeyglossa. - G. Stomosis sp. 1. Abbreviations: bs - basisternum; cv 0 - cervical organ; cx - coxa; fs - furcasternum; hd - starting point of head; 1 cv sc1- lateral cervical sc1erite; p b - precoxal bridge; p cv sci - posterior cervical sclerite; proes - proepisternum; pres - presternum; proem - proepimeron; pprn - postpronotum; sec sci - secondmy sclerite. Scales 0.1 mm.

with the cervical organ, which probably has a articulates laterally with the propleuron (PI. SB). sensory function. Laterillly of the cervical organ I shall begin with the description of the basi­ lies the lateral cervical sc1erite and posterior to it sternum, because it is phylogenetically most the narrow posterior cervical sclerite, which informative for the Milichiidae. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 37

A developed, which means that the area between the fore coxae is almost completely sc1erotised, and the basisternum extends anteriorly and laterally to join the propleuron (PI. 8C). Precoxal bridges are not neces­ sarily homologous, because there are different ways in which the bridge may be constructed (see below). In the Milichiidae all three main types ofbasis tern a exist (Speight 1969). In my opinion, the basisternum is small in the stem-species pattern of the B Milichiidae, because there are small basisterna in the stem-specics pattern of the Acartophthalmidae and Car­ nidae. In the stem-species pattern of the Chloropidae there is a precoxal bridge. This is certainly autapomorphic because there cannot be a pre coxal bridge in the stem-species pattern of 1: the Milichiidae. This is because the II two groups of the Milichiidae in which a precoxal bridge occurs, Litometopa / and Desmometopa saguaro, belong to

PI. 9. A-B. Adult, female. - A. Desl11ol11etopa l11-nigrlll11 (after Sabrosky the Madizinae. In other Desmometopa 1987a). - B. Acartophthalll1l1s nigrin!ls (Acartophthalmidae) (after J. F. species, in the stem-species pattern of McAlpine 1987). Abbreviations: aneps s - anepisternal seta; dc s - the Madizinae, and in the stem-species dorsocentral seta; keps s - katepisternal seta; M - media; npl s - notopleural seta; orb s - orbital seta; pa s - postalar seta; poc s - postocellar seta; pprn pattern of its sister-group, the Milichi­ s - postpronotal seta; prs s - presutural seta; prse s - prescutellar seta; R­ inae, there is a basiliform basisternum radius; sa s - supraalar seta; Sc - subcosta; scut s - seutellar seta. (Fig. 7). The precoxal bridge therefore has to be a novelty in Litometopa ahd in Desmometopa saguaro (see below) Basisternum (C20) and is convergent with the precoxal bridge present Three main shapes of basis tern urn were recog­ in the Chloropidae. nised by Speight (1969) in . (1) The The stem-species pattern of the Milichiidae, shape may be basilifOllli, which means that the with a small and roughly rectangular basisternum, area between the fore coxae is mostly sc1erotised is represented in Aldrichiomyza, genus nov. 1, but the sc1erite lies free in the cervical membrane Phyllomyza, Stomosis, and Xenophyllomyza (PI. (PI. 8A, B). (2) It may be small ("reduced" in 8E, G). In Neophyllomyza + Paramyia it is secon­ Speight), which means that the sc1erite occupies darily reduced to a nan-ow triangle (Fig. 7, PI. 8F). only a small portion of the area between the fore A large, V -shaped (basilifOllli) basisternum is found coxae. Consequently this area is mostly membra­ apomorphically in the Milichiinae + Madizinae neous (PI. 8E-G). (3) A precoxal bridge is (PI. 8A, B). In Desmometopa saguaro there is 38 Irina Brake ENT. SCAND. SUPPL. 57 (2000) secondarily a pre coxal bridge, formed by a con­ from the results of Speight (1969), who mentions nection between basisternum and propleuron, but that the presternum is evanescent in the Milichiinae. the sclerites are not fused (PI. 8D). In Litometopa A further difference between my observations and there is a pre coxal bridge, too, but in this case Speight's results is that he found a precoxal bridge basisternum and propleuron are fused (PI. 8C). not only in some Desmometopa species but also in The unfused precoxal bridge in Desmometopa Leptometopa and some Milichiella and Milichia saguaro could be an early stage of the fused bridge species. With the exception of Desmometopa, I inLitometopa. This would mean thatLitometopa is could not confirm these results. One explanation the sister-group of Desmometopa saguaro. The for our differing conclusions could be that Speight apomorphy for D. saguaro + Litometopa would be was using dry material in these cases. Even in the an unfused pre coxal bridge, which secondarily alcohol material that I studied, it was often very fuses inLitometopa. There are in fact no apomorphic difficult to decide whether there is a pre coxal characters for Desmometopa, which would argue bridge ornot, because the posterior cervical sclerite, against this hypothesis (see section 5.2). However, which articulates with the propleuron, is positioned though there are indications that Desmometopa is close to the basisternum and fonus a kind of paraphyletic, this still needs to be checked thorough­ functional precoxal bridge, which in my opinion is ly. Until this has been done, I will maintain the not homologous with a true precoxal bridge. In monophyly of the genus. The development of a other cases, the sclerites of the basisternum and pre coxal bridge strengthens the prothorax and could propleuron are joined in some specimens but there be an adaptation for predaceous behaviour in is a small cleft between them in other specimens. In Litometopa and Desmometopa saguaro. The fore these cases I decided against calling it a precoxal legs, which are raptorial in appearance, suggest bridge. Another explanation for our different results that these species may be predaceous, but un­ could be that the species that I studied are not the fortunately nothing is known about their behaviour. same as those examined by Speight, who does not mention species names. Apart from the strengthening of the pro thorax, for example as an adaptation to a prey-catching Proepimeron (C21) behaviour, the function of different shapes of the In the stem-species pattern of the Chloropidae basisternum in Milichiidae is, as yet, unclear. family-group there is no proepimeral seta. In genus According to Speight (1969) the universal presence nov. 1, Microsimus, and Phyllomyza, however, of the basisternum in Acalyptratae, whatever its there is a strong proepimeral seta, and inAldrichio­ shape, is presumably due to its function as the site myza, Neophyllomyza, Xenophyllomyza, and for attachment of leg musculature. Secondly, the several Paramyia species there is a nonnal to weak major characteristics of prosternal shape appear to seta. As a result it must be assumed that the depend primarily upon the role ofthe prosternum in presence of a proepimeral seta is an apomorphy for strengthening and supporting the anterior part of the the Phyllomyzinae and that it was secondarily lost thorax. in Stomosis.

Between basisternum and insertion of the fore coxae, there are secondary sclerites in species with Postpronotum (C22) a V -shaped basisternum and in Phyllomyza (PI. In the stem-species pattern of the Chloropidae 8B, D, E). Anterior to the basisternum there is the family-group there is one postpronotal seta, but in small triangular presternum. According to my Eusiphona, Microsimus, andPholeomyia there are observations, the presternum is weakly developed 2 such setae. The presence of more than one in Stomosis and Neophyllomyza, which is different postpronotal setae probably evolved three times. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 39

4.3.2. Mesothorax probably not homologous.

Only the setae on the anepisternum and Katepisternum (C24) katepisternum are phylogenetically important for In the stem-species pattern of the Chloropidae the Milichiidae (PI. 9A). family-group there is one katepisternal seta (PI. 9B). Within the Milichiidae, 2 setae in the stem­ Anepisternum (C23) species ofA ldrich iomyza + Xenophyllomyza and 2 The anepisternum is bare in the stem-species or 3 setae in Eusiphona, in the Milichia speciosa­ pattern of the Milichiidae. In the stem-species group, in Pholeomyia, and in Ulia have evolved pattern of the Chloropidae, there are no distinct convergently. It is possible that 2 or 3 katepisternal setae but only Shoti hairs on the anepisternum setae are an apomorphy for stem-species H (Fig. according to Andersson (1977). In the Acartoph­ 7). However, ifthat is the case then the number of thalmidae and Carnidae there are one or more setae katepisternal setae must have been secondarily and several hairs on the posterior margin of the reduced to one several times: (1) In Eusiphona, anepisternum (PI. 9B). there are 3 katepisternal setae in Eusiphona sp. I, In my opinion, one or more setae and several but in the otherEusiphona species there is only one hairs on the posterior margin of the anepisternum katepisternal seta. The presence of 3 setae is are part of the stem-species pattern ofthe Chloro­ probably the plesiomorphic character state for pidae family-group, because they are present in the Eusiphona, because at least in the length of the Acartophthalmidae and Carnidae. In the stem­ proboscis Eusiphona sp. I shows a less derived species of the Milichiidae and the Chloropidae character state than the other Eusiphona species, these setae and hairs were lost, and the anepisternum which have an extremely long proboscis. Eusiphona is seeondarily covered with short hairs in the sp. I could therefore be the sister-group of the Chloropidae (Fig. 7). I assume that the hairs in the other Eusiphona species. (2) The presence of just Chloropidae developed secondarily because they one katepisternal seta in Milichiella lacteipennis is are shorter and probably of a different kind (pubes­ probably derived, because there are more katepi­ cence) than the hairs in the Acartophthalmidae and sternal setae in all other Milichiella speeies. (3) Carnidae. The presence of one seta inMilichia distinctipennis Within the Milichiidae, there are apomorphically could be derived, too, because in the M distincti­ 3-4 strong setae on the posterior margin of the pennis-group, which comprises 4 species, there is anepisternum in Pholeomyia. In some Eusiphona one species with 2 katepisternal setae. Since the species there are a few weaker setae on the anepi­ number of katepisternal setae varies within the sternum in some specimens. Miliehiinae, the point of origin of more than one Sabrosky (1955) was of the opinion that the seta can only be determined after the relationships presence of anepisternal setae and a eostal noteh in within the subfamily have been resolved by means Eusiphona cooperi indicated the close relationship of other characters. between Eusiphona and Pholeomyia, with E. coo­ peri connecting the two genera. I have not seen E. 4.3.3. Wings cooperi but Sabrosky stated "mesopleuron along posterior margin with several strong, conspicuous The wing venation in the Milichiidae (PI. IDA) bristles and a few coarse setae." In Pholeomyia consists of the costa (C), rudimentary subcosta there are always 3 or 4 strong setae and no other (Se), radius (RI, R2 + 3, and R4 + 5), media (MI), smaller setae, and so the setation on the anepi­ anterior cubitus (CuAI and CuA2), and anal vein sternum in Pholeomyia and Eusiphona cooperi is (A I). Anteriorly at the base of the wing there is the 40 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A B C R2+3 ~~-~~~~====~ R4+5 r-m M1

dm-eu ,,'

",~, bm-eu H ,~",""""" , CuA1 C

H

F

en G H I

"i~~" M1 A 1 '\ \ CuA2"', H

PI. 10. A-J. Wing. - A. Phyllomyza securicornis. - B. Stomosis sp. 1. - C. Paramyia sp. 1. - D. Aldrichiomyza longirostris. - E. Litometopa ?glabriJi'ons. - F. Leptometopa latipes. - G. Desmometopa sordida. - H. Pholeomyia sp. \, female. - 1. Ulia poecilogastra, female. - J. Milichiella lacteipennis, female (after Sabrosky 1987a). Abbreviations: A - anal vein; a 1- anal lobe; bm-cu - basal medial-cubital crossvein; C - costa; c b - costal break; c n - costal notch; cell cup - posterior cubital cell; CuA anterior cubitus; dm-cu - discal medial-cubital crossvein; h - humeral crossvein; M - media; R - radius; r-m - radial-medial crossvein; Sc - subcosta. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny ofMilichiidae 41 humeral crossvein (h); between R4 + 5 and MI In the stem-species pattem of the Chloropidae there is the radial-medial crossvein (r-m), and family-group the costal vein extends to MI. In the between MI and CuAI there is the basal medial­ Milichiidae the costa convergently extends only to cubital crossvein (bm-cu) as well as the discal R4 + 5 in Paramyia and in the stem-species of medial-cubital crossvein (dm-cu). At the base of Aldrichiomyza + Xenophyllomyza (PI. IOC, D). the anal vein there are the posterior cubital cell (cell cup, "anal cell") and the anal lobe. The distal Radius and media (C27-28) margin of the posterior cubital cell is formed by In the stem-species pattem of the Milichiidae, CuA2. R4 + 5 and MI are parallel or only slightly con­ verging or diverging (PI. lOA, B, E-G), and this is Costa (C2S-26) the case in the stem-species pattern of the In the Schizophora, 2 breaks in the costa were Chloropidae family-group too: in the stem-species independently developed or lost several times (J. pattem of the Chloropidae, R4 + 5 and MI seem to F. McAlpine 1989). The humeral break lies shortly be parallel (NatiShuk 1987). They are also parallel distal to the humeral crossvein, and the subcostal in the Acartophthalmidae (PI. 9B). Fmihermore, a break occurs where the subcosta joins the costa. In parallel R4 + 5 and MI is the plesiomorphic con­ the Milichiidae and Camidae both costal breaks dition in the Camidae (present in Hemeromyia and exist and the second break lies proximal to the Neomeoneurites). Diverging veins are synapomor­ insertion ofR 1 because the subcosta is reduced. In phic for Meoneura and Camus (Fig. 2). the Chloropidae only the subcostal break is present In the Milichiidae R4 + 5 and MI are strongly and in the Acartophthalmidae only the humeral converging in Enigmilichia, Eusiphona, Milichia break. In my opinion, 2 costal breaks are part of the distinctipennis, Milichiella, and Ulia (PI. IOI-J). stem-species pattem of the Chloropidae family­ According to the cladogram (Fig. 7), converging group. The humeral break in the Chloropidae and R4 + 5 and MI are an autapomorphy for the the subcostal break in the Acartophthalmidae have Milichiinae, and parallel veins evolved secondarily been lost secondarily. in stem-species I [Milichia speciosa-group + Pholeomyia] (PI. 1OH). In Paramyia and converg­ The functional significance of the costal breaks is ently in Aldrichiomyza + Xenophyllomyza, R4 + 5 not clear. It may be an adaptation to the folding of and MI are diverging (PI. IOC, D). the wing in the puparium (Hering 1937) and/or to In Paramyia and Xenophyllomyza crossvein the skill to fly. dm-cu is convergently lost (PI. 1~C). However, in Within Milichiidae the subcostal break has Xenophyllomyza a vestigial anterior end of the apomorphically developed into a notch in stem­ crossvein is present. species H (PI. 1OH-J). This notch can be quite deep in some Pholeomyia and Milichia species. The A general trend in the evolution ofthe dipteran wing section proximal to the notch is developed into a is the concentration of veins at the anterior margin lappet, and the costa apically of the notch runs of the wing (costalisation) and the reduction ofveins along the distal side of the notch. In Eusiphona the at the posterior margin (Hennig 1973). The costal notch is secondarily lost. However, there is costalisation of the wings developed several times one species, Eusiphona cooperi, which has a notch independently, probably because offlight mechanics. at the subcostal break. It is possible that the notch In Paramyia and Aldrichiomyza + Xenophyllomyza was lost2xwithinEusiphona, because in Eusiphona the costalisation has gone very far, because (1) in all sp. 1, which probably branches off earlier within 3 genera the costa is reduced posterior to R4 + 5, (2) the genus (see section 4.3.2) there is no notch. in Paramyia and Xenophyllomyza crossvein dm-eu 42 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

is missing, (3) especially in Aldrichiomyza and 4.3.4. Legs Xenophyllomyza R2 + 3 and R4 + 5 are very close to the costa, and (4) in Paramyia Ml and euAI are There are a few phylogenetically informative weakly developed. characters on the femur, tibia, and tarsus. In Litometopa and some Desmometopa species (Sabrosky 1983), the fore legs are raptorial in Anal vein and posterior cubital cell (C29-30) appearance (see section 4.3.1), with ventral spines The anal vein (AI) ends directly distal to the on femur, elongated coxa and femur, and in posterior cubital cell in all Milichiidae. Al and Litometopa also with a striking colour pattern. CuA2 can be well developed or weak. Well devel­ This shape of the fore legs probably evolved oped A 1 and CuA2 are probably the plesiomorphic convergently in Litometopa and within Desmome­ character state, because A 1 and CuA2 are also well topa, but could also be a synapomorphy for Lito­ developed in the Acartophthalmidae and Carnidae. metopa and partofDesmometopa (see section 5.2). In both families the anal vein ends shortly before the wing margin. In the stem-species of the Femur (C31) Chloropidae, Al and CuA2 are lost apomorphically. On the hind leg of stem-species F, there is In the stem-species pattern of the Milichiidae apomorphically a long seta anteroventrally in the and of the Chloropidae family-group CuA2 is distal half of the femur. bulging (PI. lOG). In the stem-species ofthe Mili­ On the middle femur of the males of genus nov. chiinae, however, CuA2 meets Al in an acute 1, there is a femoral organ anterodorsally and angle (PI. IOH-J). Convergently with the Mili­ shortly proximal to the middle. It consists of a field chiinae, an angle between CuA2 and A 1 is present of irregularly-placed tiny setulae of probably sen­ in several Leptometopa species (PI. IOF). sory function (PI. lIF, G). Similar-looking femoral In the stem-species pattern of the Chloropidae organs on the middle legs of males are typical for family-group, the shape of the wing is oval. How­ the subfamily Oscinellinae of the Chloropidae but ever, in the Milichiidae, in stem-species H, the anal do not belong to the stem-species pattern of the lobe is apomorphically well developed and the Chloropidae according to Nartshuk (1987) and apex ofthe wing is narrow, in males more so than Andersson (1977) (Fig. 1). I have therefore not in females. This gives the wing a more triangular included this character in the data matrix. However, shape (PI. 10H-J). The condition in Enigmilichia since there is a femoral organ in genus nov. 1, it is intermediate. InXenophyllomyza the anal lobe is may be possible that the femoral organ is a also convergently well developed, but the apex of synapomorphy for the Milichiidae and Chloropidae the wing is not narrower than in the stem-species and was secondarily reduced several times. pattern of the Milichiidae.

According to J. F. McAlpine & Munroe (1968) a Tibia (C32) large anal 10 be and a triangular wing indicate species In several species of Leptometopa, the hind that swarm and complete the initial stages of mating tibia is greatly enlarged and flat in males (PI. lIE), in flight. A large anal lobe and nalTOW wing apex is and has a slight dorsal ridge in females (PI. 11 C, thought to improve the insects ability to hover and D). This means that either (1) these characters may manoeuvre with the precision required for successful be apomorphic for Leptometopa, and have been aerial coupling. The shape of the wings ofMilichiinae reduced in the L. niveipennis-group which has cOlTespond to this hypothesis, since swarming is normal hind tibiae, or (2) they may have evolved known for several species (see section 4.2.3 and within the genus, or (3) the genus is not mono­ 4.4.1 ). phyletic (see section 5.2). Convergently with ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 43

PI. 11. A-B. Madiza glabra. ~ A. Hind leg, posterior view. ~ B. Tibia, posterior view. ~ C-E. Leptolllelopa lalipes, hind leg, anterior view. ~ C-D. Female. ~C. Femur and tibia. ~ D. Distal tip of tibia. ~ E. Male, femur and tibia. ~ F-H. Genus nov. I sp. I. ~ F-G. Middle femur with femoral organ, anterodorsal view. ~ H. First tarsomere of hind leg, antcrovcntral view. Abbreviations: d r - dorsal ridge; £2(3) - middle (hind) femur; ti3 - hind tibia; ti org - tibial organ. Scales 0.1 mm. 44 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Tarsus (C33) In the Phyllomyzinae, there is apomorphically a comb-like row of setae posteroventrally on the first tarsomere of the hind leg (Fig. 8). In the stem­ species of Aldrichiomyza + Xenophyllomyza this comb is lost, and in genus nov. 1 sp. 1 there are several rows of about 3 setae each anteroventrally along the first tarsomere. These rows are arranged in a slightly diagonal direction (PI. I1H). The setae in the row ofPhyllomyza securicornis and Neophyllomyza acyglossa look like normal setae, but in Paramyia sp. 1, Stomosis sp. 1, and genus nov. 1 sp. 1 the tips of the setae are flattened. For an evaluation of the difference between setae with sharp tips and setae with flattened tips, a comparative study with the SEM is needed. I studied this character in only one species of each Fig. 8. Phyllomyza sp. 1, first tarsomere of hind leg, anterior genus with the SEM, and so cannot generalise from view. Scale 0.1 mm. the results.

4.4. Abdomen Leptometopa, an enlarged hind tibia has evolved in males of Desll1ometopa leptometopoides. For convenience, the abdomen is divided into In Madiza there is a narrow longitudinal tibial preabdomen (segments 1 to 5) and postabdomen organ at the middle of the posterior side ofthe hind (segments 6 to 11). The sc1erites ofthe preabdomen tibia of males and females (PI. llA, B). It consists are not modified by the fOlmation of the genitalia. of a long and narrow depression, densely covered The postabdomen ofmale Diptera is fmiher divided with short hairs. In addition to the tibial organ, into the pregenital segments (segments 6 to 8) and males of Madiza glabra have abdominal glands the male genitalia (segments 9 to 11). The post­ which probably produce pheromones. It would be abdomen of female Diptera forms the ovipositor. interesting to know with which behaviour the tibial organ and the abdominal glands are connected. 4.4.1. Male preabdomen (C34-36) Tibial organs on the hind legs of males and females are an apomorphy for the Chloropidae. In the stem-species pattem of the Milichiidae They resemble the organs of Madiza in structure, and ofthe Chloropidae family-group, tergites 2 to and are supposed to be either sensory areas or 5 (T2-5) are brownish microtomentose or shining secretory in function (Sabrosky 1987b). Tibial in males and females. However, in males of most organs may either be a synapomorphy for the species of the genera Milichia, Milichiella, Milichiidae + Chloropidae, and were then reduced Pholeomyia, and Ulia, T2-5 are partly or completely several times within the Milichiidae and within the silvery microtomentose. In these genera and in Chloropidae, or they may have evolved conver­ Eusiphona, the lateral margins ofthe tergites of the gently in the Chloropidae and in Madiza. I prefer males are strongly bent under the abdomen, and in the latter hypothesis, because it is more parsimo­ some Milichiella, Pholeomyia, and Ulia species nious. there is even a crease in the tergites at the site ofthe ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 45

A S4

H T6 c~~.~

o o S8

I

PI. 12. A-J. Male. - A. Stomosis sp. 1, sternites 4 and 5, ventral view. - B-D. Sternite 5, ventral view. - B. Phylloll1yza securicol'11is. - C. Milichia mYl'mecophila. - D. Pholeomyia sp. 1; E. Stomosis sp. 1, abdomen, lateral view. - F-J. Stemites 6-8. - F. Stolllosis vittata, posterior view. - G. Apotl'opina IOl1gepilosa (Chloropidae), lateral view. - H. Hemerol11yia sp. 1 (Carnidae), posterior view. - L Phyllomyza secul'icol'l1is, posterior view. - J. Desmometopa sOl'dida, anterior view. Abbreviations: dp - distiphallus; gen - genitalia; S - stemite; T - tergite; spir - spiracle. Scales 0.1 mm. 46 Irina Brake ENT. SCAND. SUPPL. 57 (2000) bend. In this case, the dorsal side of the abdomen species ofthese genera, the membrane between T5 is flat, so that the silvery surface reflects light and the horse-shoe shaped S7/8 is partly or optimally. The light-reflecting abdomen could be completely sclerotised or the lateral and distal an adaptation for swarming behaviour, so that margins ofT5 are bent ventrally. In addition, S5 is males swarming in sunlight can be seen from a considerably sclerotised and enlarged in these long distance. These swarms have been observed species (PI. 12C, D), so that the genitalia are in Milichiafumicostata, the only species ofthe M. protected by a strong chitinous ring. At present it distinctipennis-group with silvery microtomentose is not possible to decide whether these characters abdomen (Deeming 1981), and in several Milichi­ evolved once or several times. ella species (Mik 1887, IllingwOlih 1923 and 1929, The shape of the male abdomen, with silvery Malloch 1934, Duda 1935b, Cuthbertson 1936, microtomentum, flat abdomen, ventroflexion of the Hardy & Delfinado 1980, Deeming 1998). There is genitalia, and maybe also the protection of the one swarming species, Milichiella lacteipennis, genitalia by a chinitous ring could be an adaptation which does not have a silvery microtomentose to the swanning behaviour in sunlight. Females can abdomen. Instead, the abdomen is highly polished recognise the swann from a distance and visit it for and shining, and also reflects the sunlight as I finding a mate. Swanning is especially useful for myself have observed. species with scattered or thinly occupied larval The silvery micro tomentum on the tergites could habitats, for which mate finding is otherwise difficult be an apomorphy for stem-species H, in which case (Downes 1969). it must have been secondarily lost in several species, for example in Milichiella lacteipennis. Other In the stem-species pattern of the Milichiidae possibilities are that the silvery microtomentum is and ofthe Chloropidae family-group, S5 is roughly an apomorphy for a smaller group of genera within rectangular and about as long as S4. In the the Milichiinae, or that it evolved independently Milichiinae and the Madizinae, S5 is generally as within the Milichia distinctipennis-group, the long as broad or longer than broad. In the Milichia speciosa-group, Milichiella, P holeomyia, Phyllomyzinae, S5 is generally slightly shorter and Ulia. The latter hypothesis is quite improbable than broad (PI. 12B). In stem-species F [Stomosis in my opinion. I believe that the silvery micro­ + (Aldrichiomyza + Xenophyllomyza)] , S5 is tomentum evolved only once, either in stem-species apomorphically strongly reduced in size so that it H or within the Milichiinae after the branching off is much shorter than S4 or absent. This size of Enigmilichia and Eusiphona, because in all the reduction of S5 leaves a membraneous pocket in genera except for these two there are species with front of the genitalia, in which the distiphallus, a silvery microtomentose abdomen. which is very large in these genera, can be stored. Strongly bent tergites are either a novelty for the In Stomosis, S5 is divided into a narrow anterior Milichiinae or they evolved after the branching off and a narrow posterior part. The anterior part is of Enigmilichia. From the description of Enig­ bent behind S4 and covered with setulae, which milichia, I suppose that the tergites are not bent in may be part of some kind of gland (PI. 12A). this genus, but I have not studied males of Enigmilichia for myself. 4.4.2. Male pregenital segments In the genera Eusiphona, Milichia, Milichiella, Pholeomyia and Ulia, sternite 7/8 (S7/8, see below) In all the male genitalia are rotated and the male genitalia are more or less strongly through 3600 (hypopygium circumversum). This folded ventrally under the abdomen, so that they rotation occurs near segment 8 and results in a are barely visible from the dorsal side. In some shifting and often a reduction of the sclerites of ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 47 segments 6 to 8. S6-8 shift to the left and S7 and S8 ring with S7/8. often fuse to form a single synsternite lying more Within the Milichiidae, S7/8 is narrow in most or less dorsally (Crampton 1941). T6 may be species (PI. 121), but can be wider in Stoll1osis, reduced in size or lost, and T7-8 are absent. In all Madiza, and Neophyllomyza (PI. 12E). In some male Diptera the abdomen has only 7 spiracles. Phyllomyza species slits can be discerned which The 6th and 7th spiracles are either in the sclerotised possibly separate S7 from S8. These slits could area of synsternite S7/8 (PI. 121), directly next to thus indicate that this sclerite originated from two S7/8 in an only lightly sclerotised area (PI. 121), or fused sclerites. The plesiomorphic condition ofS6, in the membrane. in which it is fused to the left side of S7/8, is In the stem-species pattern of the Chloropidae represented in ?Xenophyllomyza sp. 1 and in family-group, T6 is present but narrow. This Stomosis (PI. 12F). In Milichiella sp. 2 and assumption is based on the presence of T6 in the Phyllomyza, S6 convergently joins S7/8 on each stem-species pattern of the Carnidae (Wheeler side (PI. 121). In all other Milichiidae that I studied, 1994) and of the Milichiidae. Within the Milichi­ S6 is absent. idae, T6 is present only in Madiza glabra and Stomosis (PI. 12E) and must have been lost in all 4.4.3. Male genitalia other genera. In addition, T6 is lost in the Chloro­ pidae (Andersson 1977) and in the Acartoph­ The homologisation of structures in the male thalmidae (PI. 20A). genitalia of Diptera has been a matter for con­ In the stem-species pattern of the Chloropidae siderable controversy: on the one hand, the homo­ family-group, as represented in the stem-species logy with structures in other insects is unclear, and pattern of the Carnidae (Neomeoneurites and on the other hand the character transformation Hemeromyia) and present in most Schizophora, within Diptera is disputed. Basically, the hypotheses the pregeni tal segments are strongly asymmetrical, differ in the homologisation of the saddle-shaped with S6-8 lying closely joined on the left side (PI. dorsal sclerite (epandrium or periandrium), and its 12H). This is the plesiomorphic condition, which ventral appendages, the surstyli. The main theories is caused by the circumversion ofthe male genitalia. are (1 )the epandrialhypothesis ofCrampton (1936), An apomorphy of the Acaliophthalmidae + Chillcott(1958) and Hennig (1976); (2) theperiand­ (Milichiidae + Chloropidae) (stem-species A) is rial hypothesis of Griffiths (1972, 1981, 1991); the formation of a symmetrical, dorsal S7/8 (PIs. and (3) the revised epandrial hypothesis of Cum­ 12F, G, I, 1; 20A). Similar but convergently evolved ming, Sinclair & Wood(1995). The third hypothesis forms ofsynsternites can be found, among others, is based on the first hypothesis. Because the in the . In the Tethinidae and , discussion in Cumming, Sinclair & Wood is convin­ S7/8 is symmetrical as well, but it is fused to the cing and the descriptions and illustrations are readily large T6. The presence of a synsternite S7/8 could comprehensible, I will follow the revised epandrial be an indication that the Tethinidae/Canacidae hypothesis in this study even though it has been complex is the sister-group of the Acartophthal­ criticised by some authors (Griffiths 1996, Zat­ midae + (Milichiidae + Chloropidae) (see section warnicki 1996). 6.1). However, further studies are needed to support According to Cumming, Sinclair & Wood this hypothesis. (1995), the male genitalia in Schizophora consist In the Milichiidae and Chloropidae (Andersson of the dorsal saddle-shaped epandrium, which 1977), S6 is plesiomorphically fused to the left evolved from T9, and the ventral hypandrium, side of synsternite S7/8 (PI. 12F, G). In the which evolved from a fusion of S9 and the Acartophthalmidae, S6 apomorphically forms a gonocoxites (PI. 13A). The tips of the U-shaped 48 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A ep

sep sci

ep sep sci D

ej ap~

phap

E ej~hYp \

Pi. 13. A-E. Male genitalia, lateral view. - A. DeslI1ometopa sordida (distiphallus not fully extended). - B. Litometopa ?glabriji

dph

PI. 14. A-D. Male genitalia, lateral view. - A. Stoll1osis vittala. - B. Aldrichiomyza longirostris (distiphallus not fully extended). - C. Paramyia sp. I. - D. Neophyl/omyza sp. I (distiphallus macerated). Abbreviations: bph - basiphallus; ce - cerci; dph - distiphallus; ej ap - ejaculatory apodeme; ep - epandrium; hyp - hypandrium; phap - phallapodeme; preg­ pregonitc; sep scl- subepandrial sclerite; sur - surstylus. Scales 0.1 mm. membrane of the dorsal part of segment 9. The ovipositor at copulation. paired cerci belong to the 11 th segment; between Inside the epandrium the base of the phallus, them lies the anus. The surstyli are secondary which is inserted into the female vagina at appendages ofthe epandrium. They hold the female copulation, lies at rest. The phallus consists of the 50 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

aedeagus fused with the parameres and is divided the surstylus is sharp or very narrow (PI. 20A, C). into basi- and distiphallus. At its base the phallus Within the Milichiidae this character state is present articulates with the rod-like phallapodeme, which in Desmometopa, EnigmilicJzia, Litometopa, the is thought to have originated from a longitudinal Milichia speciosa-group, and Pholeomyia (PIs. invagination of the inner wall of the hypandrium. l8B, 19B, C). The surstylus is apically rounded Laterally the basiphallus is encircled by the convergently in stem-species E, Eusiphona, Ma­ postgonites (PI. l6D, 20A, B). Anterior to the diza, Leptometopa, and Milichia distinctipennis postgonites are the pregonites, which are connected (Fig. 9; PIs. 13D, 14A-D, l8C). Milichiella is with the hypandrium anteriorly. Both postgonites characterised by a slightly spoon-shaped surstylus and pregonites probably evolved from gonopods, (PIs. l5B, 16B), and in Ulia the surstylus has 2 but pregonites were newly formed within the lappets. In Phyllomyza (PI. 16C) and gen. nov. 1 Schizophora. The phallus is moved by the muscles the surstylus is deeply notched at the middle. This ofthe phallapodeme; post- and pregonites prevent may be an apomorphy for stem-species C (Fig. 9), a lateral movement. From the phallus the ejaculatory but needs to be checked for Costalima and Micro­ duct leads to the sperm pump with the ejaculatory simus, which have not been dissected. In Stomosis apodeme, which are situated shOlily anterior to the there is a small appendage on the anterior side of epandrium inside the abdomen (PI. l3B). The the surstylus in some species (PI. 14A). In other ejaculatory apodeme was originally a part of the species there is a larger appendage (PI. 15F). aedeagus. The sperm pump consists of a membra­ The margins of the surstylus are more or less neous sac, which is squeezed in order to press smooth; only in Madiza is the distal margin densely sperm into the female vagina. dentate, and in Enigmilichia (Deeming 1981, Fig. 8) and Eusiphona (PI. l8C) the posterior margin is Epandrium evenly dentate. I do not know whether the dentation In the Milichiidae, the epandrium is mostly in Enigmilichia and Eusiphona is homologous, saddle-shaped and bears few large setae. A more because I have not studied the genitalia of Enig­ spherical epandrium is typical for Phyllomyza and milichia myself. gen. nov. 1 and may have evolved in stem-species C. However, this character needs to be checked for Hypandrium and subepandrial sclerite Costalima and Microsimus, which have not been In the stem-species pattern of the Chloropidae dissected. family-group, as represented in the Acartophthal­ In the stem-species pattern of the Milichiidae, midae, Carnidae, and Milichiidae, the hypandrium the posterodorsal part of the epandrium slopes is U-shaped and the subepandrial sclerite forms a markedly down to the cerci (PIs. l4A-B, l6C). In single plate, connecting the arms ofthe hypandrium the Milichiinae and Madizinae, this slope is only anteriorly (PI. 17D). slight or even absent (PI. l3A, B, D). In the stem-species pattern ofthe Chloropidae, the hypandrium is probably fused anteriorlyinto a Surstyli (C37-38) ring according to Andersson (1977) and N artshuk In the stem-species pattern of the Milichiidae (1987). However, according to both these authors, and of the Chloropidae family-group, the surstyli it is also possible that the ring-shaped hypandrium are not fused with the epandrium (PI. l5B). How­ evolved within the Chloropidae. The stem-species ever, such a fusion occurs convergently in Neo­ pattern condition of the subepandrial sclerite has phyllomyza and Stomosis (PIs. l4A, D; l5E, F). not been studied. It either forms a single plate, or In the stem-species pattern of the Chloropidae apomorphically it forms 2 lateral rods leading family-group (present in all 4 families), the tip of from the articulation of the hypandrium with the ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 51

PI. 15. A-F. Male genitalia. - A. Phyllomyza securicornis, lateral view. - B. Milichiella lacteipennis, posterolateral view. - C. Ma(liza glabra, phallus (distiphallUs not fully extended), lateral view. -D. Stomosis sp. 1, anteriorviewofhypandrial complex (seen frol11 the direction of the thick arrow in F). - E. Neophyllomyza sp. 1, posterolateral view. - F. Stomosis sp. I, lateral view. Abbreviations: ce - cerci; dph - distiphallus; ep - epandrium; hyp - hypandriul11; phap - phallapodeme; phap scl- phallapodemic sclerite; preg - pregonite; sur - surstylus. Scales 0.1 mm.

epandrium to the inner side of the surstyli. nected anteriorly by a narrow sc1erotised band. Within the Milichiidae, in Stomosis, the hypan­ If there is a ring-shaped hypandrium present in drium is fused into a ring (Pl. 17B), and the sub­ the stem-species pattern of the Chloropidae, as epandrial sc1erite fOllllS 2 rods which are con- suggested by Andersson and Nartshuk, then it is 52 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A ce

sur c

ep--'l sep sci I

/) ~;/) c / PI. 16. A-C. Male genitalia, lateral view. - A. Ulia poecilogastra. - B. Milichiel!a laeteipennis. - C. Phyl!omyza sp. 1. - D. Apotropina ?jilscipleuris (Chloropidae), hypandrial complex, lateral view. Abbreviations: ce - cerci; dph - distiphallus; ej ap - ejaculatory apodcme; cp - epandrium; hyp - hypandrium; pg - postgonite; phap - phallapodemc; preg - pregonite; sep scl- subepandrial sclerite; sur - surstylus. Scales 0.1 mm. possible that this holds true for the Milichiidae too. mean though that a reduction of the ring has taken If this is the case, the shape of the hypandrial place four times (Fig. 9). It is more parsimonious complex as in Stomosis would represent the stem­ to assume that in the stem-species pattem of the species pattem of the Milichiidae. This would Milichiidae the anns of the hypandrium are free ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 53

A B sur

ep

ep

ph sep sci phap sci I hyp phap preg hyp phap sci phap

o

sep sci

preg

phap sci hyp

PI. 17. A-D. Hypandrial complex, ventral view. - A. Phyllomyza secliricornis. - B. Stomosis villa/a. -- C. Paramyia sp. 1. - D. Neopi1yllomyza acyg!ossa. Abbreviations: bph - basiphallus; ce - cerci; ep - epandriull1; hypo. hypandriull1; ph­ phallus; phap - phallapodeme; phap scl - phallapodemic sclerite; preg - pregonite; sep scI - subcpandrial sclerite; sur­ surstylus. Scales 0.1 mm. 54 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

and were apomorphically fused inStomosis. In this demic sclerite evolved from the body wall anterior case the ring-shaped hypandrium evolved conver­ to the phallus and is an apomorphy for the Chloro­ gently in Stomosis and in the Chloropidae. pidae family-group (Griffiths 1972). Within the Chloropidae family-group there occur different In my opinion there is possibly a functional degrees of fusion between the phallapodemic correlation between the shape of the hypandrium sclerite, hypandrium and pregonites: and the shape of the subepandrial sc1erite. A In the Camidae, the phallapodemic sc1erite is connection between both tips of the hypandrium is either free (Hemeromyia sp., according to Griffiths probably important for the stability of the genitalia. 1972), fused laterally with the hypandrium (Meo­ This connection is ensured either by lengthening neura sp., according to Griffiths 1972; Hemeromyia and fusion of the hypandrial arms into a ring, or by sp., according to Wheeler 1994) (PI. 20C), fused a connection ofthehypandrial arms with the anterior with the hypandrium and the bases ofthe pregonites margin of the subepandrial sc1erite. In cases where (Hemeromyia sp. 1), or absent (Neomeoneurites, a ring is present, as in Stol11osis and Apotropina according to Wheeler 1994). The Camidae are ?jilscipleuris (Chloropidae), the subepandrial sc1erite therefore not suitable for an outgroup comparison can be split into two rods, which may only be ofthis character so long as the ground pattem ofthe connected by a narrow band. This hypothesis has to genitalia of this family has not been established. be checked for Chloropidae. In the Acartophthalmidae, a fusion of the phall­ apodemic sclerite with the hypandrium and the Pregonites and postgonites (C39) bases ofthe pregonites occurs, without a reduction In the stem-species pattem of the Chloropidae, of the median part of the hypandrium. According Acartophthalmidae and Camidae, pregonites as to Andersson (1977), the phallapodemic sclerite in well as postgonites are present (PIs. l6D, 20B). the Acartophthalmidae is not fused with the An apomorphy for the Milichiidae is the absence hypandrium, but only linked via membranes ofpostgonites (Griffiths 1972), so that the base of anteriorly to the hypandrium and laterally to the the phallus is enveloped by the pregonites. The discrete pregonites. Based on my own studies, I pregonites are fused with the hypandrium and have disagree with Andersson's conclusion, but agree from three to many setulae (PI. 17B-D). with Griffiths' (1972) opinion that there actually is Within the , the postgonites are ab­ a fusion. sent only in the Phoroidea, in some , In the Chloropidae there is no fusion between and in most Pipunculidae (Cumming, Sinclair & the phallapodemic sclerite, pregonites and hypan­ Wood 1995). These groups are celiainlynot closely drium. related to the Milichiidae, as they do not belong to Within the Milichiidae there is no fusion between the Schizophora. The loss of the postgonites has the phallapodemic sclerite, pregonites and hypan­ thus evolved convergently in these groups and in drium in Neophyllomyza, Paramyia, and Phyllo- the Milichiidae. I -er-tyIUijilwmyza (PI. 17 A, C, D). In Aldrichiomyza, Stomosis, andXenophyllomyza, the phallapodemic Phallapodeme and phallapodemic sclerite (C40- sclerite is fused with the median part of the 41) hypandrium (PIs. 15D, I7B). In the Milichiinae In the stem-species pattem of the Chloropidae and Madizinae, the phallapodemic sclerite is fused family-group the hypandrium encircles a sclerotised with the bases ofthe pregonites and the free anterior plate. This plate is called the phallapodemic sclerite, ends ofthe hypandrial anns. The median pati ofthe because it is fused with the posterior part of the hypandrial arc is reduced (PI. 18A~D). phallapodeme (Andersson 1977). The phallapo- Since there are different states of fusion between ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 55

Phyllomyzinae Madizinae Milichiinae I I Q) ....: co C\l ~ 91 ~ ~ ~ I U 0 - :> ~ C\l ~ C\l ~ Q) C\l .g ~ o m c E: § ~_~ ~ ~ C\l ~ 't ~ ~ ~ Q) (/) o Q) ~ 2 .9?]? .9? 0 'c: '0.. c.. co :::l C Q) (9 ~~!!ii3i~~~I~ I I I I I I ~ sujstYIU~ sp~on-~hap~d I dlstiphallus with two stripes tdistiphallus a slender I tube with hairs

surstylus rounded apically distiPha~ith JostJior ,J, surst~lus rou~dedl apically sclfrotise~ stripe I I' I I sursytYluSwith two lappets distip~allu~t with ~o stripes ~

phap sci fused Iwith surstylus rounded apically p med;" "rt o 'r

surstylus rounded apically I surstylus notched distiphallus sock-shaped

distiphallus a slender tube with widened middle I distiphallus with posterior sclerotised stripe

phap sci fused with preg and hyp I phap sci fused with preg and hyp

phap sci not fused with hyp Ciis!iphallusst\6rf& •glabrous - surstylus with narrow tip B - distiphallus flexible, coiled, and pubescent =-____--=--= ____---1 - distiphallus without sclerotisations ~~--~--~,~ A

Fig. 9. Phylogenetic tree of the Milichiidae with a depiction of the transformation of 4 characters of the male genitalia (C38 shape of surstylus, C40 extent of fusion between phallapodemic sclerite, pregonites, and hypandrium, C42 shape of distiphalIus, C43 sclerotisations on distiphaIlus). Shaded hatchmarks indicate character changes that either rcverse or evolved at least 2x on the tree; black hatchmarks indicate changes that do not reverse and evolved only once; white hatchmarks indicate plesiomorphic character states; reductions are marked with a darker shading. Abbreviations: hyp - hypandriu1l1, phap scl- phallapodemic sclerite, preg - pregonite. 56 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A

ep

ep

, hyp a phap sci I phap sci ~Phap

c D preg

hyp a

preg

phap sci phap I phap sci phap dph PI. 18. A-C. Male genitalia in posleroventral view and hypandrial complex in ventral view. - A. Pholeomyia sp. I. - B. Milichia myrmecophila (after Brake 1999). - C. Eusiphona sp. I. - D. Madiza glabra, hypandrial complex, dorsal view. Abbreviations: ce - cerci; dph - distiphallns; ep - epandrium; hyp a - hypandrial arms; phap - phallapodeme; phap scl­ phallapodemic sclerite; preg - pregonite; sur - surstylus. Scales 0.1 mm. the phallapodemic sclerite, hypandrium, and assess. Either the phallapodemic sclerite is free in pregonites in each family, the stem-species pattern the stem-species pattern and fusions occurred of the Chloropidae family-group is difficult to several times convergently, or the fusion in the ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 57

PI. 19. A-B. Desmometopa sordida, male genitalia. - A. Lateral view. - B. Ventral view. - C. Litometopa ? glabrifi'ol1s, male genitalia, lateral view. - D. Litometopa ? glabrifi'olls, female ovipositor, lateral view. - E-F. Desmometopa /11- Iligrum, ovipositor, lateral view. - E. Secondary ovipositor everted. - F. Secondary ovipositor inverted. Abbreviations: bph - basiphallus; ce - cerci; dph - distiphallus; ep - epandrium; g 0 - genital opening; r b s - ring of barbed spines; S - sternite; sba - subanal plate; sur - surstylus; T - tergite. Scales 0.1 mm

Acartophthalmidae and part of the Carnidae or reversed several times convergently. represents the stem-species pattern and was changed In my opinion there is (either plesiomorphically 58 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A B T5

S5---T"':

dph ~~"ii-- sep sci

Tl<;----T--- ce

c phap D

sur

PI. 20. A-B. Acartophthalmus nigrinus (Acartophthalmidae), male genitalia (after J. F. McAlpine 1987). - A. Lateral view. - B. Ventral view. - C. Hemeromyia sp. (Carnidae), male genitalia, ventral view (after Wheeler 1994). - D. Desmometopa sordida, male reproductive system, ventral view. Abbreviations: ace gl- accessory gland; ce - cerci; dph­ distiphallus; ej ap - ejaculatory apodeme; ej d - ejaculatory duct; ep - epandrium; hyp - hypandrium; pg - postgonite; phap - phallapodeme; phap scl - phallapodemic sclerite; preg - pregonite; S - stemite; sep scl - subepandrial sclerite; sur - surstylus; T - tergite; test - testis; vas def - vas deferens. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 59 or apomorphically) no fusion between the phall­ shaped" sac projecting anteriorly (PI. 14A-D). apodemic sc1erite, pregonites and hypandrium in The distiphalli within Stol11osis are of varying the stem-species pattern of the Milichiidae + shapes and apomorphically have special kinds of Chloropidae (Fig. 9). As a consequence it must be spines (PI. 14A). In Paramyia sp. 1, the tip of the assumed that this condition represents the character distiphallus has simple spines (PI. 14C). In state in the stem-species pattern ofthe Milichiidae. Aldrichiomyza, Neophyllol11yza, and Xenophyllo­ In this case, the fusion between the phallapodemic myza, there is posterolaterally a sc1erotised stripe sc1erite and the middle of the hypandrium is an at the base of the distiphallus on each side (PI. 14B, apomorphy for stem-species F. The fusion between D). These stripes evolved convergently in the phallapodemic sc1erite, pregonites, and hypand­ Aldrichiomyza + Xenophyllomyza and in Neo­ rial anns, combined with a reduction ofthe median phyllomyza. part of the hypandrium, is an apomorphy for the Milichiinae + Madizinae (stem-species G). Cerci In the stem-species pattern of the Chloropidae Phallus (C42-43) family-group the cerci are not fused. In the In the stem-species pattern of the Chloropidae Chloropidae the cerci are fused posteromedially. family-group, as represented in the Acartoph­ In the Acartophthalmidae there is a kind of narTOW thalmidae and Carnidae, the membraneous disti­ lip on the outer margin ofthe cerci, which is neither phallus is long, flexible, coiled, and pubescent (PI. sc1erotised nor covered with setae. 20A). An apomorphy for the Milichiidae + Chloro­ In the Milichiidae the cerci can vary in size. In pidae is a short and glabrous distiphallus. Desmometopa and Litometopa (PIs. 13A, B, 19A­ Within the Milichiidae, this short and glabrous C) they are nearly as large as the epandrium in distiphallus is present in Eusiphona, the Milichia lateral view, while they are very small in Lepto­ distinctipen nis-group, Milichiella, P holeomyia, and metopa and Neophyllomyza among others (PIs. Ulia, as well as insome species ofPhyllol11yza (PIs. 13D, 14D). Setae can be found mainly on the outer l3E; 15A; 16A, B; 18A, C). In Phyllomyza sp. 1, margin, one or 2 very large ones being apical (PI. the phallus is slender and secondarily long again, 14A). The cerci can be sc1erotised to a varying and is only slightly sc1erotised at its ventral base extent. However, in species with lightly sc1erotised (PI. 16C). In the Milichia speciosa-group, the cerci the ventral margin is often more strongly distiphalli are slender tubes with fine hairs (PI. sc1erotised. Several Milichiidae show some kind 18B). of lip on the outer margin of the cerci, which is The phalli of the Madizinae are velY similar to neither sc1erotised nor covered with setae. In each other. The distiphallus apomorphically fOllllS Desmometopa, Litometopa, Phyllomyza, and Sto­ a slender tube, which is slightly widened in the mosis, this lip is particularly strongly developed middle (PIs. 13A, B, D; 15C; 19C). At the base of (PIs. l3A, B, 14A, 15A, 19A, B). In many genera the distiphallus there is posteriorly a sc1erotised the lip is scarcely or not at all visible (PI. 15B, E). stripe, which can also be found in Ulia and in some The lips on the cerci could be an apomorphy for species of Milichia, which both belong to the the Milichiidae. In this case the lip was secondarily Milichiinae. It is therefore possible that this stripe reduced several times within the family. But the is plesiomorphic for the Madizinae and evolved in character is difficult to evaluate, because the lip is the stem-species of the Milichiinae + Madizinae often narrow or difficult to see. Among these (Fig. 9). narrow forms in particular, differences between In stem-species E, the distiphallus is apomor­ species of the same genus occur. For the exact phi cally modified into a wide and often "sock- evaluation ofthis character, the stem-species pattern 60 Irina Brake ENT. SCAND. SUPPL. 57 (2000) A B c

••• 0 C'.,' ~. 'J': .>

S7 I r--""'\--- s ba

D F

S7~

E sci strip (S8) H7

T8

sba

ce

PI. 21. A-F. Female ovipositor. -A-Co Desmometopa m-nigrum.-A Dorsal view. - B. Ventral view. -Co Lateral view. - D-E. Milichia sp. 1. - D. Dorsal view. - E. Ventral view. - F-G. Lilometopa ?glabrifi·ons. - F. Tip of ovipositor, lateral view. - G. Sternite 6. Abbreviations: ce - cerci; S - sternite; sba - subanal plate; sci strip - sclerotised strip; sec ovp - secondmy ovipositor; spa - supra-anal plate; T - tergite. Scales 0.1 111111. ENT. SCAND. SUPPL. 57 (2000) Phylogeny ofMilichiidac 61 of each genus has to be studied first. need to bc studied to confirm these hyp()lh,~scs. Variation in the genitalia at genus-level In thc Chloropidac, the testes arc ovoid or The number of different shapes of the male pyriform in thc Oscinellinae and clongak and genitalia within each genus varies. Within tubular in the Chloropinac. The vasa defCrelll iii ilrc Aldrichiomyza, Desmometopa, Litometopa, not pigmented and are not fused in mosl species Neophyllomyza, Paramyia, and Phyllomyza, the (Adams & Mulla 1967, Nartshuk 1997). male genitalia resemble each other closely in all In the Camidae, thc tcstcs are pyrifOlm, wil hi hc the species examined by me and by other authors. large part at the distal tip, and the vasa defercill iii The close similarity between the genitalia of Des­ are neither pigmented nor fused (Meoneura lIlld mometopa and Litometopa even suppOlis the hypo­ Hemeromyia). thesis that Litometopa belongs to Desmometopa In the future, when more information becomes (see section 5.2). In the genera Leptometopa and available, the shape of the intcrnal male repro­ Stomosis, there are two different shapes ofsurstylus ductive system may be useful for the differentiation in each. In Milichiella, the length of the surstyli of groups of genera. and the setation on the pregonites vary, and in Pholeomyia there are various shapes ofepandrium 4.4.5. Female ovipositor (C44-52) and surstyli. In the genus Milichia most male genitalia are very similar to each other. Only the The ovipositor of acalyptrate flies consists of genitalia of Milichia distinctipennis and probably r segments 6 to 8 and paired cerci, the latter being the whole of the M. distinctFpennis-group are appendages of segment 11. Inserted between different. This group probably does not belong to segment 8 and the cerci is another section, the the genus Milichia (see section 5.3). sclerites of which have not yet been satisfactorily The shape of the genitalia could be a diagnostic homologised. These sclerites are called S9 and T9 character for genus determination and an apo- by Matsuda (1976), hypoproct and epiproct (con- morphy of the species-groups, because it is quite sisting of elements from segments 9 and 10) by J. constant in all the known species-groups. Genera F. McAlpine (1981), or sub- and supra-anal plate with several shapes of genitalia should be checked by Andersson (1977). I am using the terms sub­ for poly- or paraphyly. and supra-anal plate, because they describe the relative positions ofthe sclerites and do not suggest 4.4.4. Male reproductive system any homologisation with the sclerites ofa particular segment. The female genital opening lies behind The male reproductive system of the Diptera S8, and the anal opening between subanal plate and consists of paired testes, paired vasa deferentia, cerci. paired accessory glands and an unpaired ejaculatory In the stem-species pattern of the Chloropidae duct. The testes are orange-brown because of a family-group and ofthe Milichiidae, the ovipositor pigment in the outer epithelium. The male repro­ is long and retractable. I suggest that all the sclerites ductive system can only be dissected in fresh of the ovipositor were originally complete, and material, and consequently information is scarce. that divisions or losses of sclerites have occulTed In the Milichiidae, the testes are egg-shaped or several times independently. I have come to this elongate (Desmol71etopa) (PI. 20D). The vasa conclusion because in all 4 families of the Chloro­ deferentia are also orange-brown and are basally pidae family-group there are species with complete fused. The pigmentation and fusion of the vasa sclerites as well as species with longitudinally deferentia could be apomorphies for the Madizinae divided or absent sclerites. It is more probable to or even for the Milichiidae. However, more species assume that sclerites were divided or lost several 62 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Phyllomyzinae Madizinae Milichiinae I I C\l ....; C\l C\l ....; OJ ~ I ~ C\l 0.. 9' E: ,.9 .~ ....; :> V) C\l E: .S2 ~ 0 C\l ::J 0 E: (])~-f:C\l~ c ~ .~ -s.. E: ,.9 .~ § v :lg .§ 'S., .~ V) ..c:: .Q V) ~ V) E: ..c:: 0 0.. ..c:: :::J .i:l .S2 0.. E: 0 .g c V) e E: ~Et_-%~~ (,) 0 ~ c:: Q) 0 -s.. (]) C\l ,.9 ~ <.9 () ~ 5: <: Q C/) ~ <:( ~ ~ Gj LD ~ ~ :§

S8 'Iongi'tudin'ally divid~d, I ring absent, sec ovp absent, X S8 ~bs~nt, ri~g a~sen(i sec ovp absent, X 1 S8 complete,U ring absent, S8 ?, rin~sent,1 sec ovp a~sent, ~, tsec ovp absent, 1 S810ngitudlnally divided, I ~S8110~9' divided, ring L---"r_ln_g--,present, sec ovp jbseT X absent, sec ovpabsenr

S8 halfway longitudinally divided 'S8 absent, ring present, : I secovp absent, X ~ S8 longitudinally divided F

S8 longitudinally divided, ring present,secovp absent; X c E H

ring <;Ibsen!' - S8 forming two strips - secondary ovipositor present - ovipositor short - spa bare G - cerci short

ring €Ibsen! I . S8 longitudinally divided 0

- S8 complete - ring between segment 7 and 8 - secondary ovipositor absent - ovipositor long - supra-anal plate with one pair of setae - cerci long

Fig. 10. Phylogenetic tree of the Milichiidae with a depiction of the transformation of6 characters of the ovipositor (C47 shape of S8, C48 ring of barbed spines, C49 secondary ovipositor, C50 length of ovipositor, C51 setae on supra-anal plate, C52length of cerci). Shaded hatchmarks indicate character changes that eitherreverse or evolved at least 2x on the tree; white hatchmarks indicate plesiomorphiccharacter states; reductions are marked with a darker shading. "X" marks the combination ofthe three characters: ovipositor long, supra-anal plate with one pair of setae, and cerci long. Abbreviations: S - sternite, sec ovp - secondary ovipositor, spa - supra-anal plate. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 63

times, rather than that they are secondarily undi­ species pattem of the Milichiidae. Distally there vided or secondarily present. The same holds true are 2 weakly sclerotised strips, each with a row of for the evolution of a ring of barbed spines between setulae. These strips originate ventrolaterally and segments 7 and 8: this ring can be found in some meet dorsally atthe functional tip ofthe ovipositor. species in each family ofthe Chloropidae family­ Between these strips, which represent S8, lies the group as well as in several other families of genital opening. When the secondary ovipositor is acalyptrate Diptera, and must either have evolved retracted (PI. 19D, F), only the basal part is tumed repeatedly or have been lost several times inde­ over, with the barbed spines now directed posteri­ pendently. I suggest that the ring of barbed spines orly (PI. 2lF). In addition to the secondary ovi­ is pati ofthe stem-species pattem ofthe Milichiidae positor, Desmometopa, Litometopa, and Pholeo­ and of the stem-species pattem of the Chloropidae myia are characterised by a bare supra-anal plate family-group and that it was lost several times and short, round cerci. This is in contrast to the independently. stem-species pattem of the Milichiidae and of the Within the Milichiidae, the plesiomorphic state Chloropidae family-group, where there is one pair of the ovipositor sclerites (all sclerites undivided) of setulae on the supra-anal plate and the cerci are is present only in gen. nov. 1 and in Leptometopa elongate. 1- niveipennis. More frequently, S8 is modified or The close similarity of the complex-structured ...--~., .. absent, and in addition S7, T7, or T8 are sometimes secondary ovipositor in Desmometopa, Litometopa divided (PIs. 2lA-E, 22A-L). The plesiomorphic and Pholeomyia suggests that it is a homologous state of the ring of small barbed spines between structure and evolved only once. As a consequence, segments 7 and 8 is only present in Enigmilichia the secondary ovipositor must have evolved in the dimorphica, Eusiphona sp. 1, some Leptometopa stem-species of the Madizinae + Milichiinae and species, and Madiza glabra (PI. 22G, H). In the was reduced once or 2x in the Madizinae and Phyllomyzinae, in the Leptol1letopa niveipennis­ several times in the Milichiinae (Fig. 10). group, and in several Milichiinae the ring is lost A secondary ovipositor occurs only in the (Fig. 10). In contrast to this, the barbed spines are Milichiidae, but similar, though not homologous, strongly developed in Desmometopa, Litometopa, structures can be found in e.g. the Agromyzidae and Pholeomyia. (Spencer 1987) and the Drosophilidae (Grimaldi In Desmometopa ,Litometopa, andPholeomyia, 1987). In the Agromyzidae, segment 7 is modified the ovipositor is short, with T6-7 and S6-7 into a non-retractable oviscapt into which the undivided, S8 specialised, and T8 considerably following segments can be retracted. The membrane shortened (PIs. 19D, F; 21A-C, F). The shape ofS6 between segments 7 and 8 bears anteriorly pointed is triangular in Desmometopa and Pholeomyia (PI. denticles of different sizes and shapes, which are 2lB) and has a median projection on the anterior used to tear open plant tissue. S8 is developed into margin in Litometopa (PI. 21 G). Inside segment 7, paired "egg guides". females have a specialised structure which is described here for the first time and which is called The function of the secondary ovipositor in the secondary ovipositor. At oviposition the secon­ Desmometopa ,Litometopa, and Pholeoll1yia is not dary ovipositor is everted telescope-like and then known. The resemblance between the secondary ovipositor and the ovipositor of plant-mining lies between S7 and the sub anal plate (PI. 19E). Agromyzidae may indicate that the females of these Basally the structure consists of a broad mem­ milichiid genera lay their eggs into relatively fresh braneous ring with many anteriorly-directed barbed plant material, which decomposes afterwards. spines. This ring probably originated from the ring Several Desmometopa species have been bred from of spines between segments 7 and 8 in the stem- decomposing plant material and larvae of Pholeo- 64 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

A B c o

E F I

J

D

T7 87

T8 T8 88 I

Pl. 22. A-L. Female ovipositor. - A-B. Phyllomyza secliricornis. - A. Dorsal view. - B. Ventral view. - C-D. Milichia sp. 2. - C. Dorsal view. - D. Ventral view. - E-F. Leptometopa latipes. - E. Dorsal view. - F. Ventral view. - G­ H. Madiza glabra (from Sabrosky 1987a). - G. Lateral view. - H. Dorsal view. - 1-1. Neophyllomyza sp. 3. - 1. Dorsal view. - J. Ventral view. - K-L. Sto1l1osis vittata. - K. Dorsal view. - L. Ventral view. Abbreviations: ce - cerci; S­ sternite; sba - subanal plate; spa - supra-anal plate; T - tergite. Scales 0.1 mm. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 65

myia have been found in waste chambers of leaf­ ovaries and unite to form the common oviduct (PI. cutting ants. The biology ofLitol11etopa and the egg­ 23A). The common oviduct opens into the vagina. laying behaviour of Milichiidae is completely un­ Dorsally two spermathecal ducts and the ducts of known. The strong, barbed spines may indicate that the paired accessory glands open on the genital the females hook up on the substrate. This would papilla into the vagina (Pis. 23B, C, 26A). In the make it easier for the female to push the ovipositor into more solid substrates. The short, and with its schizophoran stem-species pattern there are three complete sclerites strong ovipositor ofthe species in spermathecae (Downes 1968, Hennig 1973, J. F. question indicates a more solid substrate for egg­ McAlpine 1989). However, the number of sper­ laying as well. mathecae was frequently reduced to two. The spermathecae consist of a sclerotised capsule In the Milichia speciosa-group there are species sun-ounded by epithelial gland cells (Kotrba 1993) with a short ovipositor (Milichia speciosa, Milichia (PI. 26C). In the ventral wall ofthe vagina, opposite sp. 1) which is very similar to the ovipositor of the genital papilla, there is an unpaired ventral Pholeomyia: T6-8 and S6-7 are undivided, the receptacle in several acalyptrate families (Sturte­ anterior margin of S6 is convex, S8 is divided into vant 1925-1926, Kotrba 1993). Like the sperma­ narrow strips, the pair of setulae on the supra-anal thecae, the ventral receptacle functions as a spellll plate is absent, and the cerci are short and round. reservoir. When an egg passes through the vagina, However, the ovipositor of these Milichia species the micropyle is placed next to the entrance ofthe does not have the secondary ovipositor (PI. 21D, receptacle and insemination takes place. In Mi1i­ E). There are also Milichia species with a long chiidae the ventral receptacle is followed posteriorly ovipositor, for example Milichia sp. 2 (PI. 22C-D). by an epithelial pad of unknown function. I believe that a bare supra-anal plate and short cerci are pati of the character complex ofthe secondary Spermathecal ducts (C53) ovipositor, and so the shape ofthe ovipositor in the In the stem-species pattern of the Chloropidae Milichia speciosa-group is probably a transitional family-group, as represented in the stem-species stage between an ovipositor with secondary pattern ofthe Carnidae, the spermathecal ducts are ovipositor and one in which it has been lost. ShOli. This situation corresponds to the stem-species InLeptometopa the degree ofsclerite subdivision pattern of the Acalyptratae. in the ovipositor differs strongly within the genus. Within the Carnidae, ShOli spermathecal ducts L. niveipennis has undivided sclerites, whereas L. are present in Neomeoneurites (Hennig 1972) and latipes and L. rufifrons, for example, have Hemeromyia (PI. 26B, C). In Meoneura, an longitudinally divided T7, T8, and S8, as well as elongation and tangling of the ducts occurs more or less completely divided S6 and S7 (PI. apomorphically at the base of the spermathecal 22E, F). The Leptometopa niveipennis-group and ducts within the muscular sheath ofthe vagina (PI. the L. latipes-group, to which L. rufifrons belongs, 26A) (Kotrba, pel's. comm.; Kotrba & Brake, manu­ differ in other characters as well (Papp 1984), and script in preparation). the monophyly of the genus needs to be checked In the stem-species of the (Milichiidae + (see section 5.2). Chloropidae) + Acartophthalmidae (stem-species A), the spermathecal ducts are apomorphically 4.4.6. Female reproductive system extremely elongated and are rolled up together distally into one longish spool-like coil (Pis. 23C, In the Schizophora, the abdomen of sexually 24A). In situ this spermathecal duct coil lies mature females is taken up mainly with the paired ventrally at the base of the ovaries (PI. 23A). ovaries, filled with eggs. Short oviducts start at the Basally the spermathecal ducts are wider and 66 Irina Brake ENT. SCAND. SUPPL. 57 (2000) possibly sUlTounded by muscles. In the coil the and more loosely wound, i.e. the ducts do not lie ducts and the sUlTounding epithelium are nalTower. directly next to each other. In Milichiella, there are The elongation of the spermathecal ducts in species whose spermathecal ducts represent the stem-species A is not homologous with the elonga­ stem-species pattern, as well as species with very tion in Meoneura, because (1) in Meoneura the tightly wound, spindle-like spermathecal duct coils elongation occurs within the muscular sheath of (PI. 25B). According to the drawing ofPh ole omyi a the vagina, i.e. basally, while in the other group it indecora by Stutievant (1925-1926), the coil is occurs distally; (2) the spermathecal ducts form a spindle-like in this species too, but in my dissection tangle in Meoneura instead of a coil; (3) the ofPholeomyia sp. 1 the coil was longish and spool­ phylogeny of the Carnidae is well-founded: the like. This would mean that a spindle-like genera Neomeoneurites and Hemeromyia, which spetmathecal duct coil evolved convergently within have the short ducts that are also present in other two genera. Schizophora, branch off earlier than Meoneura In my opinion it is improbable that a spindle-like (Hering 1972, Grimaldi 1997) (Fig. 2). Itis therefore spermathecal duct coil evolved 2x. It may be possible more parsimonious to assume that the elongation that Sturtevant accidentally put the wrong title to the evolved only once in Meoneura (and maybe in drawing, because his drawing is quite similar to my Camus) than that it is part of the stem-species drawing ofMilichielia lacteipennis, aspecies, which pattern of Carnidae and was secondarily reduced he examined as well. Sturtevant did not comment on 2x. the different shapes of the coil in his text. Further As mentioned above, the spetmathecal ducts dissections of other Milichiella and Pholeomyia are long and are rolled up together into one longish species, especially of Pholeomyia indecora will spool-like coil in the Acartophthalmidae (PI. 23C). solve this problem. In the stem-species pattern of the Chloropidae (present in Siphonellopsinae and Oscinellinae), The Milichiidae, Chloropidae, and ACatioph­ the spermathecal ducts are also elongated and are thalmidae are the only known families of the rolled into a coil (Oscinellinae: Sturtevant 1925- Acalyptratae with considerably elongated and 1926, Schwartz 1965, Adams & Mulla 1967, Pol­ distally coiled spermathecal ducts (Sturtevant lock 1996). However, unlike the Acartophthalmidae 1925-1926). The spermathecal duct coil is probably and Milichiidae, these coils are not longish, but are oblong in stem-species A, because this is the case apomorphically circular (PI. 25D, E). The sper­ in the Acartophthalmidae and Milichiidae. In the mathecae lie in the middle of the coil. In the Chloropidae, the coil is apomorphically circular. Chloropidae, there are species with lengthened but Very large coils are plesiomorphic, because this not coiled up spetmathecal ducts in the subfamily character state is present in the ACatiophthalmidae, Chloropinae (Sturtevant 1925-1926) (PI. 26D). most Milichiidae, and several Chloropidae. Secon­ In the stem-species pattern of the Milichiidae, dary shortening of the duct, or reduction in the size the paired spermathecal ducts are rolled up into of the coil, occulTed within the Milichiidae and one longish coil, as in the Acartophthalmidae (Stur­ Chloropidae. tevant 1925-1926) (PIs. 24A,B,E; 25A, C). Within the Milichiidae, there are only a few modifications As discussed by Lachma"nn (1994), the elongation of the spermathecal ducts. The ducts may be of of the spermathecal ducts may be connected with different lengths, i.e. the coil has a different number sperm competition after sperm transfer. The sperm ofloops, or the coils are differently wound up. In of the last copulating male, which is closest to the the stem-species of Desmometopa + Litometopa opening of the spermathecal duct, probably has the (PI. 24F, H), the coils are apomorphically smaller best chance of inseminating an egg, while the sperm ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Miliehiidae 67

B A ov v ree

______sp d

~ ace gl

epith p

vag e------ee I sp d el sp d C

ovd com

v ree

ace gl d

sp

-T-t--vag

PI. 23. A-C. Female reproductive system. - A. Schematic drawing of thc Milichiidae in situ, ventral view. - B. Leplomelopa lalipes, lateral view. - C. Acarlophlhalmus l1igril1us (Acartophthalmidae), lateral view. Abbreviations: acc gl- accessory gland; acc gl d - accessory gland duct; ce - cerci; epith p - epithelial pad; ov - ovary; ovd - oviduct; ovd com - oviductus communis; sp - spennatheca; sp d - spermathecal duct; sp d cl-spermathecal duct coil; v rec - ventral rcceptacle; vag - vagina. Scales 0.1 mm. 68 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

D pump org

sp

sp

PI. 24. A. Phyllol11yza secllricornis, spermathecal duct coil. - B-C. Leptol11etopa rufifi·ol1s. - B. Spermathecal duct coil. - C. Accessory gland. - D-E. Madizaglabra. - D. Accessory gland. - E. Spermathecal duct coil. - F-G. Deslllol11etopa sordida. - F. Sperl11athecal duel coil. - G. Accessory gland. - H. Litometopa ?glabrifi'ol1s, spennathecal duct coil. Abbreviations: pump org - pumping organ; sp - spermatheca. Scales 0.1 111m. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 69

D

B

PI. 25. A-E. Spermathecal duct coils. - A. Milichia distinctipennis (one spermatheca lost during dissection). - B. Milichiella iacteipennis. - C. Phoieoll1yia sp. 1. - D. Oscinellinae (Chloropidae). - E. Apolropina seniiis (Siphonellopsinae, Chloropidae). Abbreviations: sp - spennatheca; sp cap - spermathecal capsule. Scales 0.1 mm. 70 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

of previous copulating males is pressed back. In 26D). A small spennathecal capsule, as is present inseminated Milichiidae spermatozoa can be found in the Oscinellinae, probably represents the stem­ mainly within the ventral receptacle, but also in the species pattern of the Chloropidae because the wider basal part of the spennathecal ducts and presence of a capsule is the plesiomorphic character sometimes in the coil as well. state for the Chloropidae family-group and for the Acalyptratae. As a result, the capsule must have Spermathecae been reduced several times within the Chloropidae. In the stem-species pattern of the Chloropidae The short and slender duct at the end of the sper­ family-group, as represented in the stem-species mathecal duct is probably an apomorphy of the pattern of the Carnidae, there are sc1erotised Chloropidae (Fig. 1). sperrnathecal capsules with spherical sUlTounding In the stem-species pattern of the Milichiidae, tissue (PI. 26C). Sc1erotised spermathecal capsules there is no sc1erotised spelmathecal capsule. The are considered plesiomorphic for the Chloropidae distal end ofthe spennathecal ducts projects slightly family-group because they are present in the stem­ out ofthe coil and is surrounded by epithelial gland species pattern of the Acalyptratae. cells. There is only slight variation in the shape of Within the Carnidae, the stem-species pattern is the spermathecae within the Milichiidae: the represented in Neomeoneurites and Hemeromyia spelmathecae may be rather long (PIs. 24B, 25B) (PI. 26B, C). However, in Meoneura, the capsules or more spherical (PI. 25A). In some species the are reduced in size and can be found as small, distal end of the spennathecae is pointed (PIs. 24E, sc1erotised, pointed tips at the end of the distally 25B). In the stem-species of Desmometopa + Lito­ slightly enlarged spermathecal ducts (PI. 26A). metopa the distal end ofthe ducts is slightly enlarged In Acartophthalmus nigrinlls (Acartophthal­ (PI. 24F, H). midae), the spermathecal capsules are completely The shape of the spermathecae may be useful for reduced and the sperrnathecal ducts end with slight phylogenetic studies. However, as yet not enough ramifications in the sUlTounding tissue (PI. 23C). species have been studied, because unsclerotised In the Chloropidae, a sperrnathecal capsule is spelmathecae are very difficult to detect in preserved sometimes present and the shape of the spenna­ material. thecae differs: the tissue sUlTounding the end of the spermathecal ducts is always spherical in the Since spelmathecal capsules are present in the Siphonellopsinae and Oscinellinae that I studied, stem-species pattern of the Chloropidae family­ while it is spherical or unevenly shaped in the group and in the stem-species pattern of the Chloropinae. In the Oscinellinae the tips of the Carnidae and Chloropidae, the capsule must have sperrnathecal ducts are slightly enlarged and are been reduced in the Acartophthalmidae, in the connected to the mostly very small, lightly stem-species of the Milichiidae, and at least 2x sc1erotised spermathecal capsule via a short and within the Chloropidae. A complete reduction of slender duct (PI. 25D). In the Siphonellopsinae, the the spermathecal capsule also occurs convergently tips of the sperrnathecal ducts are also slightly in several acalyptrate families, for example in the enlarged, but there is no capsule (PI. 25E). In these Ephydridae and Fergusoninidae. cases the short and slender duct opens into a lumen surrounded by gland tissue. In the species of the Accessory glands subfamily Chloropinae that I examined, the sper- The shape of the accessory glands in the stem- mathecal ducts end without an enlargement, but species pattern ofthe Chloropidae family-group is also with a Sh01t, slender duct which opens into a uncertain, because only a few glands have been lumen within the sUlTounding gland tissue (PI. described. The reason for this deficiency is that, ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 71

PI. 26. A. Meoneura iacteipennis (Carnidae), female reproductive system, lateral view. - B. Neomeoneurites chi/ensis (Camidae), spermatheca. - C. Hemeromyia sp. I (Carnidae), spermathecae. - D. Chloropinae (Chloropidae), female reproductive system, lateral view. Abbreviations: acc gl d - accessory gland duct; ce - cerci; epith p - epithelial pad; gen pap - genital papilla; ovd com - oviductus communis; sp - spermatheca; sp cap - spermathecal capsule; sp d - spermathecal duct; v rec - ventral receptacle; vag - vagina. Scales 0.1 mm. 72 Irina Brake ENT. SCAND. SUPPL. 57 (2000) because of their small size and soft structure, In Hemeromyia the ventral receptacle does not accessory glands are difficult to demonstrate in have a stem and the lumen seems to be small. Since fresh material and nearly impossible to detect in the shape of the ventral receptacle in the stem­ dry or alcohol material. species pattern ofthe Chloropidae family-group is The accessory glands ofthe Acartophthalmidae unknown, the stem-species pattern ofthe Carnidae are unknown, and the accessory glands of the cannot be asceltained. carnid Meoneura vagans have long ducts and are The ventral receptacle in the acartophthalmid attached to the base of the ovaries. Acartophthalmus nigrinus is elongate and ventrally In the chloropid subfamilies Oscinellinae and curved, like a hook (PI. 23C). Chloropinae the accessOlY glands are long and A pocket-like ventral receptacle is an apomorphy with a short duct. for the Milichiidae + Chloropidae (StUltevant 1925- Within the Milichiidae the accessOlY glands of 1926). Within the Chloropidae this shape ofventral Leptometopa spp. and Madiza glabra have velY receptacle is present in the Oscinellinae for example short ducts, while the glands of Desmometopa (Schwartz 1965). Within the Chloropinae there are sordida andPholeomyia sp. 1, have long ducts (PI. species in which the wall near the opening of the 24C, D, G). At the base of the gland there is a club­ receptacle bulges into the lumen of the receptacle, shaped widening in the duct, which is interpreted leaving space for sperm only near the outer wall as a pump. Similar pumping organs have been (PI. 26D). In these species spenn is stored in the described for other Acalyptratae (Klostenneyer & enlarged basal part of the spennathecal ducts. I Anderson 1976, Solinas & Nuzzaci 1984, Kotrba believe that a study of the female reproductive 1993, Lachmann 1994). In Leptometopa spp. and system would contribute valuable characters for Madiza glabra the gland is quite long. In Desmo­ the phylogeny of the Chloropidae. metopa sordida and Pholeomyia sp. 1 the gland is Within the Milichiidae the ventral receptacle is shorter and has a lumen. The distal end of the pocket-like in all the species that have been studied accessory gland is attached to the base of the (PI. 23B). ovaries. The shape of the accessory glands in the stem-species pattern of the Milichiidae cannot be 4.5. Immature stages ascertained, because the shape in the stem-species pattern of the other families of the Chloropidae There is such little infolmation available about family-group is also uncertain. the morphology of the eggs, larvae, and puparia of the Milichiidae that it cannot be used for phylo­ Ventral receptacle genetic systematics. Nevertheless, I will cite all the The ventral receptacle in the stem-species pattern papers that contain morphological descriptions, as ofthe Chloropidae family-group is non-chitinised a basis for future studies. and unchambered. The exact shape of the ventral receptacle in the stem-species pattern of the Chlo­ 4.5.1. Eggs ropidae family-group is unknown, because it differs within the family-group as well as within the There is only one thorough description of the Acalyptratae. egg ofMilichiella argyratoides, by RoMcek (1995). In the Carnidae, at least two shapes of ventral In addition, there is an illustration of the egg of receptacle are represented. In Meoneura the Madiza glabra in Papp & Wheeler (1998), and I am receptacle is globular and is separated from the including SEM-photographs of the eggs of this vagina by means of a short, broad stem (PI. 26A). species (PI. 27A-C). ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 73

PI. 27. A-C. Madiza glabra, egg. - A. Lateral view. - B. Anterolateral view. - C. Posterolateral view. - D-H. Desmometopa val'ipalpis, puparium. - D. Anterior end, dorsal view. - E. Anterior spiracle. - F. Posterior end, ventral view. - G. Posterior spiracle. - H. Puparium, dorsal view. Abbreviations: a sp - anterior spiracle; an - anus; mp - micropyle; p sp - posterior spiracle. Scales 0.1 mm (except where otherwise stated). 74 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

4.5.2. Larvae and puparia position below lower eye margin. Face concave in profile, median carina low. There are descriptions of the third instar larvae Proboscis slightly elongate and geniculate. Palpi of Desmometopa spp. (Zimin 1948, Hennig 1956 small and spatulate. Stipes rod-like; lacinia and ["?Leptometopa latipes" see Papp & Wheeler ventral appendage absent. Four pseudotracheae 1998]), Leptometopa coquilletti (Hennig 1956), present; pseudotracheal ring tips rounded. and Pholeomyia comans (Moser & Neff 1971). Thorax with no differences between male and Puparia are known for Desmometopa sp. (Bohart female. Basisternum small and roughly rectangular. & Gressit 1951), Enigmilichia dimorphica Mesonotum covered with hairs. 1 postpronotal, 1 (Deeming 1981), Milichia pubescens (Deeming & presutural, 2 notopleural, 1 supra- and 2 postalar, Baez 1985), Milichiella spp. (Bohart & Gressit no intraalar, 2 postsutural dorsocentral, 1 prescu­ 1951, Roh::icek 1995),Pholeomyia comans (Moser tellar acrostichal and 2 scutellar setae. Anepi­ &Neffl971), andPhyllomyza spp. (Hennig 1937) sternum and anepimeron bare; no proepimeral, 1 (see also the summaries in Ferrar 1987, Sabrosky proepisternal, and 1 katepisternal seta. 1987a, andPapp & Wheeler 1998). I am including Wings oval in shape, hyaline, without a distinct SEM-photographs of the puparium of Desmome­ colour pattern. Two costal breaks present; second topa varipalpis (PI. 25D:-H). (;' . break proximal to insertion ofRl. Subcostareduced but distinct in basal part. Rl, R2 + 3, R4 + 5, Ml, 4.6. Stem-species pattern of Milichiidae CuA 1, CuA2, and A 1 present. Costa extending to Ml; R4 + 5 and Ml parallel. Anal vein ending Based on the results discussed in sections 4.2- directly distal to the posterior cubital cell, which is 4.4, I summarise the stem-species pattern of the well developed or weak; CuA2 bulging distad. Milichiidae as follows: Halteres white. Head with no sexual differences between male Legs rather long and slender, with irregular and female. Arista slender and shOli-pubescent. hairs or setulae. Mid tibia with apical spine. Scape very small. Pedicel short, cup-shaped, with Abdomen rather narrow, microtomentose or a row of setulae along the dorsal and outer lateral shining, with moderate-sized hairs or setulae. edge, dorsally with a strong erect and slightly Tergites large, laterally broadly enfolded. S 1-5 of outcurved seta. First flagellomere relatively small medium size. and spherical, with short pubescence. Lunule well In males T6 probably present. T7-8 absent. developed, with one pair ofsetae. Eyes ofmoderate Synsternite S7/8 symmetrical, with S6 fused to its size, rounded or slightly higher than wide. Postfrons left side. Epandrium saddle-shaped, with few large slightly longer than wide, with a distinct ocellar setae. Posterodorsal part of epandrium sloping triangle. Orbital and frontal plates distinct, with 3 down to cerci. Surstyli simple and with narrow tip, lateroclinate orbital setae and 2 medioclinate frontal not fused with epandrium. Margins of surstyli setae. Postocellar setae well developed and probably more or less smooth. Hypandrium U-shaped. crnciate. Lateral vertical setae lateroclinate, medial Postgonites absent, so that base of phallus is envel­ veliical setae medioclinate, ocellar setae latero­ oped by the pregonites. Pregonites fused with proclinate. Up to 2 medioclinate paravertical setu­ hypandrium, probably with :3 setulae. Phallapode­ lae. Postfrons with a paired row 00-4 interfrontal mic sclerite present. No fusion between phallapo­ setulae and with a small proclinate setula at anterior demic sclerite and pregonites or hypandrium. Sub­ margin between supra-antennal seta and eye epandrial sclerite fonning single plate, connecting margin. A row of postocular setulae behind the hypandrial arms anteriorly. Distiphallus short and eye. Gena with one row of setae. Vibrissa distinct, membraneous. Ejaculatory apodeme small. Cerci ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 75 well developed, not fused. • basistemum large and V-shaped (C20-2), Female ovipositor long and slender, retractile. • phallapodemic sclerite fused with bases of S6-S and T6-S probably undivided. A ring of small pregonites and hypandrium (C40-2), barbed spines between segments 7 and S. Cerci • median part ofhypandrium lost (C41-1). elongate. Ventral receptacle non-chitinised, un­ A further apomorphy of the Madizinae + Mili­ chambered, and pocket-like. Paired spennathecal chiinae is probably the presence of a secondary ducts very much elongated and rolled up together ovipositor (C49), though it must have been sub­ distally into one longish spool-like coil. No sclero­ sequently reduced several times within the Mili­ tised spennathecal capsule present. Distal ends of chiinae and the Madizinae. The following character spermathecal ducts projecting out of the coil and states, which are all cOlTelated with the presence of sUlTounded by epithelial gland cells. the secondary ovipositor in my opinion, are also Biology: Larvae saprophagous. Adults probably apomorphic for the Milichiinae + Madizinae: • S S forming two strips, each with a row of setulae kleptoparasitic. (C47-4), • ring of barbed spines between S7 and SS (C4S- 5. Phylogeny of the Milichiidae 2), • secondary ovipositor present (C49-1), In this section I will list the apomorphic charac­ • ovipositor short (C50-1), ters for each group of the Milichiidae and discuss • supra-anal plate bare (C51-1), • cerci short (C52-1). problematic taxa. The evaluation of the characters has been given in the preceding morphological sections. The phylogenetic tree ofthe Milichiidae, 5.1. Phyllomyzinae which includes stem-species A to I, is shown in Fig. 3. I propose the new subfamily PhyIlomyzinae for The Milichiidae are a monophyletic family (Fig. the genera Aldrichiomyza, Costalima, gen. nov. 1, 3). Apomorphies for the Milichiidae are: Microsimus, Neophyllomyza, Paramyia, Phyllo­ • upper orbital seta lateroclinate (C 1-1), myza, Stomosis, andXenophyllomyza. These genera • middle orbital seta lateroclinate (C2-1), are excluded from the Madizinae because the • lower orbital seta lateroclinate (C3-1), remaining genera ofthe Madizinae are more closely • two medioclinate frontal setae present (C4-1), related to the Milichiinae than to those forming the • proclinate setula between supra-antennal seta PhyIlomyzinae. The PhyIlomyzinae are characteris­ and eye margin present (C6-1), ed by the following apomorphies: • a pair of setulae present on the lunule (C7-1), • the two medial pseudo tracheae converging at • four pseudotracheae (C 16-1), tips (CI7-1), • postgonites lost (C39-1). • proepimeral seta present (C21-1), • first tarsomere of hind leg with a row of setae The Milichiidae can be divided into 3 sub­ posteroventrally (C33-1), groups: Phyllomyzinaen. subfam., Madizinae, and • the ring of barbed spines between segments 7 Milichiinae. The Phyllomyzinae is the first to and S in females lost (C4S-1). branch off. The Madizinae and Milichiinae are sister-groups. A feature hypothesised to be an apomorphy of Apomorphies for the Madizinae + Milichiinae stem-species C is the notched surstylus (C3S-5). are: This character state is present in Phyllomyza and • upper orbital seta medio-reclinate (CI-2), gen. nov. 1, but needs to be checked for Costalima • postocellar setae parallel to slightly converging and Microsimus. (C5-2), I tentatively suggest that the monotypic genera 76 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Costalima and Microsimus are closely related to This division ofT7 may either have evolved once Phyllomyza, based on the enlargement of the first in stem-species E and have been reversed in Aldri­ flagellomere in males (C9-1). An apomorphy for chiomyza, or it evolved convergently in Neophyllo­ Microsimus + Phyllomyza could be the presence of myza + Paramyia, in Stomosis, and inXenophyllo­ long and thick palpi (C14-2). To verify this myza. relationship, the labella, male genitalia and ovi­ An apomorphy for the stem-species of Neo­ positor of Microsimus and Costa lim a need to be phyllomyza + Paramyia is the basistemum forming studied. An additionalapomorphy for stem-species a narrow triangle (C20-1). Additional apomorphies D could be mYlmecophily, because all three genera may be the shapes of S8 and T8 (C45-2, 47-2). are known to share this behaviour: Phyllomyza These last two characters are uncertain, because species have been found in the nests of Lasius and there are species with differently shaped S8 and T8 Formica ants, while Costalima was found in an in Neophyllomyza andParamyia. To support these Azteca nest and Microsimus in a Crematogaster characters, the stem-species pattem ofthe genera nest. In the light oftheirrelationship, the geographic Neophyllomyza andParamyia should be checked. distribution of these three genera is of interest. In stem-species F there is apomorphically a Phyllomyza is known from the Nearctic, Palaearctic, long seta on the hind femur (C31-1), S5 in males is Afrotropical, Oriental, and Australasian-Oceanian either very short or absent (C36-1), and the phall­ Regions, but not from South America. However, apodemic sclerite is fused with the median pati of although Phyllomyza is widcly distributed, its the hypandrium (C40-1). association with ants is only known from Europe. Aldrichiomyza + Xenophyllomyza are apomor­ Microsimus and Costa lima are only known from phi cally characterised by an absent upper, a reclinate South America. As all three genera are mylmeco­ middle, and a proclinate lower orbital seta (Cl-3, philous, the distribution pattem of the flies could 2-0, 3-2), by a bare lunule (C7-0), by long and well reflect the distribution ofthe ants: Formica is narrow palpi (C 14-1), by 2 katepistemal setae distributed in the N earctic and Palaearctic Regions, (C24-1), by the costa extending only to R4 + 5 and Lasius additionally in the Oriental Region. (C26-1), by diverging R4 + 5 and M1 (C27-2), by Azteca is only known from the Neotropics, but the absence of the comb on the first tarsomere of Crematogaster is worldwide in distribution (Holl­ the hind leg (C33-0), and by 2 sclerotised stripes on dobler & Wilson 1990). Neither the Phyllomyza the distiphallus (C43-2). Of these character states, flies nor the ants Formica and Lasius, with which C 14-1 (long and narrow palpi) is the only one they are associated, are found in South America. which evolved only once within the Milichiidae. However, neither are these ant genera found in the The other character states evolved more than once: Afrotropical, Australasian, and Oceanian regions. for example, C26-1 (costa extending only to R4 + The question thus arises: with which ants are the 5) and C27-2 (diverging R4 + 5 and M1) are also flies associated in these regions? present in Paramyia, and C43-2 (2 sclerotised The sister-species of stem-species C is stem­ stripes on the distiphallus) is present in Neophyllo­ species E. Stem-species E is characterised by myza. The character states which evolved con­ several apomorphic characters: long labella (C18- vergently inAldrichiomyzq + Xenophyllomyza and 1) with 2 pseudotracheae (C 16-3), apically rounded in Paramyia, C26-1 and C27-2, and the character surstyli (C38-1), and a large, "sock-shaped" which evolved convergently in Xenophyllomyza distiphallus (C42-3). Furthelmore, T7 ofthe female and Paramyia, C28-1 (crossvein dm-cu lost), are ovipositor is longitudinally divided in almost all all connected with the costalisation of the wing, the species which descended from stem-species E which took place in the stem-species of Aldri­ (C44-1). Only in Aldrichiomyza is T7 not divided. chiomyza + Xenophyllomyza and in Paramyia ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 77 convergently (see section 4.3.3). The costalisation case Desmometopa would be paraphyletic. must have evolved convergently in these two groups The sister-group relationship between Desmo­ because the sister-group relationship between metopa andLitometopa is suppOlied by two charac­ Aldrichiomyza + Xenophyllomyza and Stomosis ters: the ring tips at the distal end of the medial (stem-species F) is well supported and it is unlikely pseudotrachea in females are longer than the other that the costalisation was then reversed in Stomosis. ring tips (CI9-1), and the spelmathecal ducts are In contrast with my results, Papp (1993) wrote loosely coiled (C53-1). In addition, the male that Xenophyllomyza and Paramyia seem to be genitalia (shapes ofhypandrial complex, surstyli, related or even congeneric (he does not state why, and cerci) are very similar in Desmometopa and but it is probably because crossvein dm-cu is Litometopa (PI. 13A, B). However, Litometopa missing, C28-1) and that Aldrichiomyza and shares even more characters with the subgenus Paramyia are related because they share the costa Platophrymia of Desmometopa. Litometopa extending to R4 + 5, diverging R4 + 5 and Ml, a resembles several Platophrymia species in the very long proboscis, and because they have no shining black vibrissal angle and concave face, in vibrissa. However, Papp did not include Stomosis the cheeks being about 0.1 x the height of an eye, in and Neophyllomyza in his considerations, nor did the yellow colour of fore coxae (D. flavicoxa he make any phylogenetic analysis. According to species-group), and in the fore legs, which are my studies, which are based on many more raptorial in appearance (D. melanderi, D. meridio­ characters, there is more support for the hypothesis naZis, D. saguaro). Furthermore, the shape of the thatAldrichiomyza andXenophyllomyza are sister­ precoxal bridge in D. saguaro could be an early groups and that they are not closely related to stage of the shape ofthe pre coxal bridge found in Pal'amyia. Litometopa (see section 4.3.1). According to my studies, these character states in Platophlymia are 5.2. Madizinae all apomorphic in relation to the stem-species pattern of Desmometopa. The new ly defined subfamily Madizinae, which If Litometopa belongs to Desmometopa, then it now comprises only the genera Desmometopa, should possess the apomorphic characters of Madiza, Leptometopa, and Litometopa, is charac­ Desmometopa. The only known apomorphic terised by the shape of the distiphallus, which is character of Desmometopa is a set of interfrontal apomorphically a membraneous tube with a stripes. These, as well as the interfrontal hairs, are widened part in the middle (C42-2). In addition, not present in Litometopa. But as Sabrosky (1983) there is posteriorly a sclerotised stripe at the base pointed out, this character is problematic because of the distiphallus (C43-1), but this stripe is also there are several unnamed polished black Neo­ present in Ulia and Milichia, which belong to the tropical species that may belong to Desmometopa. Milichiinae. This character may therefore be In these unnamed species there are no interfrontal plesiomorphic for the Madizinae and may have stripes but only interfrontal hairs. According to evolved in the stem-species of the Madizinae + Sabrosky, these conditions may be degrees in the Milichiinae. reduction of the interfrontal stripes, ending with The nature ofthe relationship between Desmo­ the condition in Litometopa (without either inter­ metopa and Litometopa is unceliain. According to frontal stripes or hairs). The presence ofinterfrontal the cladistic analysis (Fig. 3), Litometopa is the stripes in Desmometopa is thus not sufficient to sister-group of Desmometopa. However, another exclude Litometopa from Desmometopa. A possibility is that Litometopa belongs to the phylogenetic study of Desmometopa and Lito­ subgenusPlatophlymia ofDesmometopa, in which metopa is needed to determine the relationship 78 Irina Brake ENT. SCAND. SUPPL. 57 (2000) between these two genera. As long as there is no 5.3. Milichiinae certain prooffor the paraphyly of Desmometopa, I will suggest that there is a sister-group relationship The subfamily Milichiinae comprises Enig­ between Desmometopa and Litol1letopa. l1lilichia, Eusiphona, the Milichia distinctipennis­ Litometopa is characterised by several apo­ group, the Milichia speciosa-group, Milichiella, morphic characters: (1) upper and lower orbital Pholeomyia, Pseudomilichia, and Ulia. The Mili­ seta lost (Cl-3, 3-3), (2) ventral side of palpi chiinae are characterised by the following apomor­ polished in females, (3) pre coxal bridge present phies: middle orbital seta reclinate (C2-0), (C20-3), and (4) males with large sclerotised sacs lower orbital seta proclinate (C3-2), within the abdomen, which may represent glands. vibrissal angle obsolescent and vibrissa above It would be interesting to see whether any of these lower margin of eye (C8-1), apomorphies are to be found withinPlatophrymia. eyes enlarged (CIO-l), The relationship between Leptometopa and frons in males nanowerthan in females (Cl1-1), Madiza, and the relationship between these two R4 + 5 and MI converging at tip (C27-1), and Desl1lometopa + Litometopa, is uncertain. distal margin of anal cell meeting anal vein in a Apomorphies for Leptometopa + Madiza could be sharp angle (C29-1). The position of the vibrissa above the lower the apically rounded surstylus (C38-1) and the loss margin ofthe eye is conelated with the enlargement ofthe secondary ovipositor (C49-0, 50-0, 51-0, 52- of the eyes, and this enlargement and the nanow 0). However, these character states are also present frons in males are probably adaptations to swarming in several other Milichiidae and may have evolved behaviour in males (see section 4.2.2 and 4.2.3). convergently in Leptometopa and Madiza. For the Enigmilichia is the first genus to branch off in present I prefer to leave the relationship between the Milichiinae. Apomorphies for stem-species H, Leptol1letopa and Madiza unresolved, with the the sister-group of Enigmilichia (Fig. 3), are possibility left open that either Leptometopa or subcostal break in the costa developed into a notch Madiza is more closely related to Desl1lometopa + (C25-1) and the triangular shape of the wing (C30- Litometopa. 1). Additional apomorphies for stem-species H There are two groups ofspecies inLeptometopa, could be a pubescent outer surface of the labella the L. niveipennis-group and the L. lalipes-group. (CI5-1) and strongly bent tergites in males (C35- TheL. latipes-group is characterised by an enlarged 1). However, the proboscis ofEnigl1lilichia has not hind tibia in males (apomorphic) and one or two been dissected, nor has the male abdomen been anepimeral setae (apomorphic). TheL. niveipennis­ studied. It is therefore not known whether these group is characterised by a setose anepistemum characters evolved in the Milichiinae before or (apomorphic) and a pair of strong prescutellar after the branching off of Enigmilichia. Further setae (apomorphic orplesiomorphic) (Papp 1984). apomorphies of stem-species H could be the The only character present in both groups is the presence of more than one katepistemal seta (C24- epistoma which is warped forward and has a I) and silvery microtomentose tergites in males triangular shape. The tip of the triangle nearly (C34-1). In that case, however, these character meets the large lunule (PI. 3B). This character state states must have been reversed in several taxa (see is expressed in Madiza in a similar way, but is not section 4.3.2 and 4.4.1). as pronounced. I cannot rule out the possibility that According to my studies, the genus Milichia is Leptometopa is not monophyletic and that Madiza para- or polyphyletic: the Milichia speciosa-group is closely related to the Leptometopa niveipennis­ is more closely related to Pholeol1lyia than to the group. Milichia distinctipennis-group. Traditionally, the ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 79 genus Milichia has been characterised only by Pseudomilichia schnusei. However, Hennig had plesiomorphic characters, for example by an entire not seen M. nigricosta. I have only seen the type­ posterior eye margin (in contrast to Milichiella and specimen ofM. nigricosta, but from the description Ulia) and a bare anepistemum (in contrast to Pho­ of P. sclmusei I conclude that both species are leomyia). Consequently it is not surprising that the valid. In my opinion, Macromilichia and Pseudo­ genus is not monophyletic. Apomorphies of stem­ milichia are both synonyms ofPh ole omyia, because species I (Milichia speciosa-group and Pholeo­ there are no good grounds for separating these taxa myia) are the loss of the setula between supra­ and because all three genera are characterised by antennal seta and eye margin (C6-0), and parallel the apomorphies of Pholeomyia. R4 + 5 and Ml (C27-0). Both character states are The relationship between the other taxa of the reversals, and so the group is not well supported. Milichiinae (Eusipholla, Milichia distillctipellllis­ The Milichia distinctipennis-group comprises group,Milichiella, and Ulia) could not be resolved M. distinctipennis, M. jumicostata, M. savanna­ in the present study. However, I believe that the ticola, and M. sylvicola. An apomorphy for the M. relationships within the Milichiinae can be resolved distinctipennis-group is that the frontal and orbital through a comparative study of the male genitalia setae in males are all reclinate except for the lower of all available species. This study should also medioclinate frontal seta. An apomorphy for the resolve the monophyly of the genera. M. speciosa-group, which comprises nearly all the other Milichia species, is the absence offrontal 5.4. Minimum age of the Phyllomyzinae, setae (C4-0) and perhaps also the tubular shape of Madizinae, and Milichiinae the distiphallus (C42-4). The relationship to theM. speciosa-group of all those Milichia species in The minimum age of the three main groups of which frontal setae are present (for example M. Milichiidae can be determined through the study ludens) should be checked. of amber inclusions. I have studied 3 Phyllomyza, Synapomorphies for the two Pholeomyia species 3 Milichiella, and 1 ?Milichia species in Dominican studied by me are the presence of 3 frontal setae amber, and 1 Phyllomyza and 1 ?Pseudodesmome­ (C4-2), of more than one postpronotal seta (C22- topa species in Baltic amber. In addition to these, 1), and on or 4 strong anepistemal setae (C23-3). Phyllomyzajaegeri and Pseudodesmometopa suc­ The generic position ofone ofthese two Pholeo­ cineum have been described from Baltic amber myia species, Pholeomyia nigricosta, has been (Hennig 1967, 1971 b) and Phyllomyza hurdi from disputed. It was described as type-species of the Mexican amber (Sabrosky 1963). genus Macromilichia by Hendel (1932). According The speciation event separating the stem-species to Hendel, Macromilichia is similar to Pseudo­ ofthe Phyllomyzinae, Madizinae, and Milichiinae milichia Becker, 1907, and Pholeomyia Bilimek, must have occuned at least 35 million years ago, 1867. In the same paper, Hendel synonymised because species which belong to the Phyllomyzinae Pseudomilichia with Pholeomyia because he con­ and species which belong to the Madizinae are vincingly argued that the character which separated represented in Baltic amber, which is 35-40 million Pseudomilichia fromPholeomyia, an emargination years old (Poinar 1992). The species which belongs ofthe posterior eye margin, was not suitable for the to the Madizinae is Pseudodesmometopa succi­ differentiation of genera. Hennig (1939) did not neum. Pseudodesmometopa may belong to the follow Hendel's synonymy, but gave no reasons stem lineage of Leptometopa + Madiza or to the for this. Instead, he synonymised Macromilichia stem lineage ofLeptometopa (Fig. 3). Pseudodes­ with Pseudomilichia and suggested that Macro­ mometopa succineum resembles the extant Lepto­ milichia nigricosta might be synonymous with metopa niveipennis, but this similarity is based on 80 Irina Brake ENT. SCAND. SUPPL. 57 (2000) plesiomorphies. The species which belong to the within the family (Nartshuk 1987). Within the Phyllomyzinae arePhyllomyzajaegeri andPhyllo­ Milichiidae, coprophagous behaviour evolved in myza sp. 6. These species differ from extantPhyllo­ the Madizinae (Desmometopa, Madiza, Lepto­ myza species only in characters that are typical for metopa) and in the Milichiinae (Milichia, Milichi­ species differentiation. Since there are twoPhyllo­ ella). However, there are always coprophagous myza species in Baltic amber, it can be assumed and saprophagous larvae in these genera. A specia­ that the genus Phyllomyza is at least 35 million lisation from saprophagous behaviour took place years old. Furthermore, the speciation events sepa­ several times in connection with myrmecophily: rating the stem-species of Phyllomyza from the larvae of Costalima live in Azteca nests, of stem-species of gen. nov. 1 and from stem-species Microsimus in Crematogaster nests, of Phyllo­ E must have occUlTed before that time. myza in nests of Lasius and Formica, and Pholeo­ In Dominican amber, which is 15-20 million myia larvae live in the nests of leaf-cutting ants. years old (Itunalde-Vinent & MacPhee 1996), The myrmecophily in Costalima, Microsimus, and there are 3 more Phyllomyza species, 3 Milichiella Phyllomyza evolved probably only once, in stem­ species, and 1 species which resembles Milichia species D, while the myrmecophily inPholeol11yia ludens. The genus Milichiella is therefore at least evolved convergently. A similar specialisation from 15 million years old. saprophagous behaviour took place in Eusiphona The milichiid genus which is most often found and some Pholeomyia larvae, which live in thc in amber is Phyllomyza. This is reminiscent ofthe nests of leaf-cutting bees. extant species, which can also be collected near The adults in the stem-species pattern of the trees. These Phyllomyza species are myrme­ Milichiidae probably fed on nectar. WithinDesmo­ cophilous on Formicinae, which have their nests in metopa, Milichia and Milichiella this behaviour or near trees. I suggest that mynnecophily evolved developed into the additional feeding on aphid in stem-species D and that consequently the stem­ honeydew. Within Mifichia it developed further species of Phyllomyza was also myrmecophilous. into being fed by ants. I believe that feeding on The fossil Phyllomyza species may have been so aphid honeydew and being fed with honeydew by frequently "caught" by the amber tree resin because ants evolved from nectar feeding because both of their myrmecophilous behaviour. honeydew and nectar consist of carbohydrates. However, the nectar-producing flowers which are 5.5. Evolutionary "trends" in the biology of the right size for small flies are not always in of the Milichiidae flower when adults are on the wing. It is therefore an advantage to be able to feed on aphid honeydew In this section I want to describe the evolution in the interim. ofmore generalised behaviours such as saprophagy In my opinion, kleptoparasitic behaviour belongs to more specialised behaviours such as myrmeco­ to the stem-species pattern of female Milichiidae phily, as it can be derived from the phylogenetic because it is present in species of all three main tree of the Milichiidae. groups (Desl11ometopa, Leptometopa, Mifichie11a, The larvae in the stem-species pattern of the Neophyllol11yza, Paramyia, Phy11ol11yza, Sto/J1osis) Milichiidae are saprophagous, because this beha­ and because it is present ii1 several Chloropidae as viour is found in most species ofthe Madizinae and well and may also belong to the stem-species Milichiinae, as well as in the stem-species pattern pattern of the Chloropidae. Kleptoparasitism pro­ of the Chloropidae according to Nartshuk (1987). vides the females with protein, which they need to The Chloropidae are commonly known as plant produce eggs, and it is thus an essential addition to miners, but this behaviour evolved several times nectar feeding. In the ancestral state, Milichiidae ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 81

AUSTRALlMYZIDAE \ surstyH fused with epandrium parameres long and strong \ ejaculatory apodemevery long

BRAULIDAE body reduced and consolidated eyes and ocelli lost sculeHum,wlngs and halteras losl claws and pulvilli modified 1 spermatheca

CARNIDAE 2 mediociinate lower fronto-orbital setae 1 interfrontal seta paravertlcal setae strong 3 postpranolal setae d abdominal membrane selu lose ejaculatory apodeme minuto C( cerci fused TETHINIDAE vibrissal angle with protuberance labella (ong, geniculate o » zJJ CANACIDAE Q lower fronto-orbital setae laterocUnate o lace protuberant m gena with upturned bristles » clypeuswide acdeagusshort 9- cerci spinose MILICHIIDAE frons wah interfrontal strips C nolchedat se break

RISIDAE first flagcllotl18re elongate face with median tubercle CendingatR4+ 5 crossvein dm·cu absent <.( sternile 8 spinulose

CRYPTOCHETIDAE larvae specialized parasites of monophlebine scales first flagellomere elongate arista reduced mouthparts reducod ,specialized <.;? terminaliaspeciallzed 1 spermatheca CHLOROPIDAE crossvein bm·cu lost cell cup lost M with peculiar inflexion scapular bristles developed

Fig. II. Phylogenctic tree ofthc Carnoidca (aftcr J. F. McAlpine 1989).

were probably attracted to predatory insects or nigrum and D. sordida. This specialisation could spiders which fed on bugs because of the defensive be the reason for the successful worldwide diffusion spray released by bugs when attacked, which is of these two species through commerce, since the used by the flies for location. WithinDesmometopa, honey bee has been distributed worldwide by man a specialisation for honey bees as the prey of as well. predatory insects or spiders evolved in D. 111- 82 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

6. Phylogenetic position ofthe Milichiidae Schizophora, for example in the Tethinidae and within the Schizophora Canacidae. However, the presence of a symmetrical S7/8 could also be an indication thatthe Tethinidael Since the position ofthe Milichiidae within the Canacidae complex is the sister-group of stem­ system of Acalyptratae is still uncertain, I will first species A (see in section 6.1). describe the sister-group relationship of the The sister-group of the (Milichiidae + Chloro­ Milichiidae. I will then work my way up to the pidae) + Acartophthalmidae is probably the Car­ family-group level. The position of the Milichiidae nidae. This group of four families was called the and related families at the superfamily level will be Chloropidae family-group by Griffiths (1972). discussed in section 6.1. According to Andersson (1977), the only valuable apomorphic character for the Chloropidae family­ The sister-group of the Milichiidae is the Chlo­ group is the presence of a phallapodemic sclerite. ropidae (Fig. 3). The following characters evolved SUPPOli for the inclusion of the Call1idae in the in their common stem-species B: • ventral appendage of stipes lost and lacinia of Chloropidae family-group is therefore rather weak. maxillae strongly reduced (C 13-1), • proboscis slightly elongate and geniculate, • an episternum without setae (C23-1), 6.1. Comparison with traditional • distiphallus short and glabrous (C42-1), classifications • pocket-like ventral receptacle. The ShOli distiphallus and the pocket-like ventral The Milichiidae, Chloropidae, and related receptacle are valuable characters because, accord­ families have been variously placed in two different ing to Griffiths (1972) and Sturtevant (1925-1926), superfamilics: and Carnoidca, or they are present only in the Milichiidae and Chloropoidea (Griffiths 1972, Hennig 1973,1. F. Chloropidae. The other characters evolved con­ McAlpine 1989). 1. r. McAlpine changed the vergently among some other Schizophora: in parti­ name Chloropoidea to Carnoidea because the oldest cular' a bare anepistel11um is widespread within family name is Carnidae. the Schizophora. An additional apomorphy could According to Griffiths (1972), the Chloropidae be the strongly converging to cruciate postocellar family-group belongs to the superfamily Tephri­ setae (C5-1). However, as the postocellar setae toidea (his prefamily Tephritoinea), which he have differing inclinations in the Acartophthal­ characterised mainly by an elongate, flexible and midae and Call1idae and since the sister-group of coiled distiphallus, bearing numerous fine cuticular the Chloropidae family-group is unknown, converg­ processes. Apart from the Chloropidae family­ ing postocellar setae may be plesiomorphic for the group, the Tephritoidea contained for example the Chloropidae family-group. , Odiniidae, Tephritidae, and Tethinidae. This superfamily in the sense of Griffiths is no The sister-group of the Milichiidae + Chloro­ longer recognised as several taxa have been pidae is the Acartophthalmidae. In their common removed to other superfamilies (D. K. McAlpine stem-species A the following characters evolved: 1985, J. F. McAlpine 1989). This means that the • spermathecal ducts elongated and rolled into a shape of the distiphallus is homoplasious. single coil distally, Hennig (1973) united the Milichiidae, Carnidae, • S7/8 symmetrical (male). and Chloropidae in the superfamily Chloropoidea, The first character is very important, because based on characters present in the Milichiidae and this situation is not known in other Schizophora. Chloropidae (spermathecal duct coil, reduced anal The second character occurs convergent! y in other vein, presence ofvibrissa, only one sclerite complex ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 83 anterior to the male genitalia). He stated that the Two families have to be excluded from the relationship between Carnidae and Milichiidae Carnoidea: the Risidae belong to the Ephydridae needed further study. Colless & McAlpine (1975) (Freidberg, Mathis & Kotrba 1998), and the expanded the Chloropoidea to include the do not belong to the Carnoidea (D. Tethinidae, Canacidae, and possibly Crypto­ K. McAlpine 1976, Wheeler 1994). It is possible chetidae, which were later placed near the Droso­ that the Tethinidae/Canacidae complex (Freidberg philoidea (D. K. McAlpine 1976). D. K. McAlpine 1995) is correctly placed in the Carnoidea because (1982) later excluded the Carnidae from the super­ these flies have asimilar habitus. However, support family Chloropoidea because the family did not for the inclusion of the Tethinidae/Canacidae agree with the Chloropoidea in several characters. complex in the Carnoidea is rather weak. Recon­ A more recent discussion of the phylogeny of struction of the stem-species pattern of the the Diptera was presented by J. F. McAlpine (1989). Tethinidae/Canacidae complex may reveal the true He placed the Milichiidae in the superfamily Car­ position of this taxon within the Schizophora. noidea, consisting ofthe families , The only genus Australimyza of the Australi­ ?, Canacidae, Carnidae, Chloropidae, myzidae has been included in the Carnidae by Cryptochetidae, Milichiidae, Risidae, and Colless & D. K. McAlpine (1991) and Grimaldi Tethinidae (fig. II). The Acartophthalmidae were (1997). However, I will neither discuss the position not included in the Carnoidea, but were placed in ofthe Australimyzidae nor the Braulidae because the . I have not studied these families. 1. F. McAlpine lists the following apomorphies J. F. McAlpine (\989) placed the Acartoph­ for the Carnoidea: thalmidae next to the in the Opomyzoidea, • arista dOl'sal, but his three arguments arc rather weak (anepi­ • vibrissa weakly differentiated, sternum with one seta ncar middle of posterior • upper fronto-orbital setae ("orbital setae") margin, S6 in male reduced, and two spennathecae). lateroclinate, In my opinion, the arguments for placing the • postocellar setae weakened, Acartophthalmidae as the sister-group of the • paravertical setae developed, • proepisternal seta developed, Milichiidac + Chloropidae (spermathecal duct coil • proepimeral seta developed, and phallapodemic sclcrite) arc much stronger. • sc weakened apically, close to R I, • 2 spermathecae. 7. Acknowledgements In my opinion, the support for the Carnoidea is rather weak because it rests mainly on characters I am especially grateful to Prof. Dr. W. Sudhaus of the chaetotaxy, which arc often variable even for supervising my doctorate studics, providing within families. Only the last character of2 sperma­ access to the facilities ofthe Institut flir Zoologic thecae is more substantial. However, it is a reduc­ of the Freie Universitiit Berlin, and for critically tion, and 2 spermathecae arc present in several reviewing thc manuscript. I am very grateful to Dr. other Schizophora fami lies (Kotrba 1993) and may A. Freidberg for acting as a referee for this papcr. very well be plesiomorphic for the Carnoidea. I thank the AG Evolutionary Biology for advice Another character which is widespread in the and discussion, and I am grateful to M. Godccke Schizophora is the dorsal arista. A proepimeral for tcchnical instruction and continuous seta is certainly not present in the stem-species encouragement. My special thanks arc due to Dr. pattern of Carnoidea, but evolved eonvergently K. Kiontkc for long discussions, hclp with tcchnical within the Milichiidae and within the Tethinidael and theoretical problcms, and for critically Canacidae complex. reviewing thc manuscript. 84 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

I thank Dr. M. Kotrba for advice and discussion, Bezzi, M. 1928. Diptera Brachycera and Athericera of the Fiji Islands based on material in the British Museum and for reviewing part of the manuscript. I thank (Natural History). British Museum (Nat. Hist.), London: Dr. H. Ulrich for reviewing part of the manuscript. viii + 220 pp. I am indebted to the NAFCiG Berlin (Nachwuchs­ Biro, L. 1899. Commensalismus bei Fliegen. - Termeszetr. Fiiz. 22: 198-204. f6rderungsgesetz) for a grant. The Agriculture and Bohart, G. E. & Gressit, J. L. 1951. Filth-inhabiting flies of Agri-Food Canada kindly allowed the use ofseveral Guam. - Bull. Bernice P. Bishop Mus. 204:1-152. figures of the Manual of Nearctic Diptera. These Brake, I. 1999. Prasaetamilichia de Meijere: a junior sub­ jective synonym ofMilichia Meigen, with a phylogenetic figures were reproduced wi th the pennission of the review of the myrmecaphila species-group (Diptera, Minister ofPublic Works and Govemment Services Milichiidae). - Tijdschr. Ent. 142:31-36. Canada, 2000. Bristowe, W. S. 1931. Notes on the biology of spiders. VII. Flies that triumph over spiders. - Ann. Mag. nat. Hist. Dr. J. C. Deeming and Dr. A. Freidberg kindly 10:471-474. invited me to study their collections, showed me Chillcott, J. G. T. 1958. The comparative morphology of the places where Milichiidae could be found, and gave male genitalia of muscoid Diptera. - Proc. 10th int. Congr. Ent. (Montreal 1956) Vol. 1:587-592. advice, for all of which I am very grateful. Coffey, M. D. 1966. Studies on the association of flies with I am indebted to Drs J. E. Chainey (NHM), P. S. dung in Southeastern Washington. - Ann. ent. Soc. Cranston (ANIC), J. C. Deeming (NMW.z), H. de Am. 59:207-218. Colless, D. H. & McAlpine, D. K. (1974) 1975.34. Diptera Jong (ZMAN), A. Freidberg (TAU), C. & H. W. (flies). Pp. 91-96 in: The insects ofAustralia. Supplement Hoffeins, J. W. Ismay, M. Kotrba (HU), W. N. 1974, Melbourne University Press, Carlton, Victoria. Mathis (USMN), B. Merz (ETH), A. L. Ozerov, G. Colless, D. H. & McAlpine, D. K. 1991. 39. Diptera. Pp. 717-786 in: The insects of Australia, Vol. 2, Melbourne O. Poinar, H.-P. Tschorsnig (SMN), and J. Ziegler University Press, Carlton, Victoria. (DEI), for the loan of material or facilitating the Collin, J. E. 1921. Description of a new genus and two new study of specimens in their collections. My special species of Cecidomyidae, and six new species of Acalyptrate Muscidae (Ephydridae and Milichiidae).­ thanks are due to Dr. M. v. Tschimhaus (UBI) for Trans. R. ent. Soc. Land. 1921:504-5 I 7. the loan of his complete alcohol collection of Crampton, G. C. 1936. Suggestions for a new interpretation Milichiidae, which has been the basis for my study, of the postabdomen in male cyclorrhaphous Diptera. - Bull. Brooklyn ent. Soc. 31:141-148, pI. VII. for giving me access to his large literature collection, Crampton, G. C. 1941. The terminal abdominal structures of and for having introduced me to this most male Diptera. - Psyche, Camb. 48:79-94. fascinating group of the Diptera. Cumming, 1. M., Sinclair, B. J. & Wood, D. M. 1995. Homology and phylogenetic implications of male genitalia in Diptera - Eremoneura. - Ent. scand. 26: 120- 8. References 15 I. Cuthbertson, A. 1936. Biological notes on some Diptera of Adams, T. S. & MulIa, M. S. 1967. The reproductive biology Southem Rhodesia. - Occ. Pap. Rhod. Mus. 5:46-63. of Hippelates col/usaI' (Diptera: Chloropidae). I. Mor­ Czemy, L. & Strobl, P. G. 1909. Spanische Dipteren. III phology of the reproductive system with notes on the Beitrag. - Verh. zool.-bot. Ges. Wien 59:121-301. physiological ageing. - Ann. ent. Soc. Am. 60: 1170- Deeming, J. C. 1981. Some new African Milichiidae (Diptera: 1176. Cyclorrhapha) having pronounced sexual dimorphism Aldrich, J. M. 1926. Descriptions of new and little-known of the frons. - Zool. 1. Linn. Soc. 71:159-169. Diptera or two-winged flies. - Proc. U.S. natn. Mus. 69 Deeming, J. C. 1998. Milichiidae and Carnidae (Diptera: (Art. 22):1-26. Cyclorrhapha) from the Arabian Peninsula. - Fauna of Aldrich, J. R. & Barros, T. M. 1995. Chemical attraction of Arabia 17:147-157. male crab spiders (Araneae, ) and klepto­ Deeming, J. C. & Baez, M. 1985. Some Milichiidae (Diptera, parasitic flies (Diptera, Milichiidae and Chloropidae). Cyclorrhapha) from the Canary Islands. - Ento­ - J. Arach. 23:212-214. mologist's man. Mag. 121:63-69. Andersson, H. 1977. Taxonomic and phylogenetic studies Donistharpe, H. S. K. 1927. The guests of British ants: their on Chloropidae (Diptera) with special reference to Old habitats and life-histories. Routledge, London. xiii + World genera. - Ent. scand., Suppl. 8:1-200. 244 pp. Becker, T. 1907. Die Dipteren-Gruppe Milichiinae. - AnnIs Downes, 1. A. 1968. Notes on the organs and processes of hist.-nat. Mus. natn. hung. 5:507-550. sperm-transfer in the lower Diptera. - Ent. Can. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 85

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Leaf-cutting ants and leaf-riding flies. Sabrosky, C. W. 1959. A revision of the genus Pholeomyia - Eco!. En!. 5:305-306. in North America (Diptera: Milichiidae). - Ann. ent. Wheeler, T. A., 1994. A new species of Neomeoneurites Soc. Am. 52:316-331. Hennig (Diptera: Carnidae) from Argentina, with a Sabrosky, C. W. 1963. A new acalyptrate fly from the discussion ofmale postabdominal strncture in the genus. 88 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

- Can. Ent. 126:435--441. Zatwarnicki, T. 1996. A new reconstruction of the origin of Wilton, D. P. 1961. Refuse containers as a source of flies in eremoneuran hypopygium and its implications for Honolulu and nearby communities. - Proc. Hawaii. classification (Insecta: Diptera). - Genus 7: 103-175. ent. Soc. 17:477-481. Zimin, L. S. 1948. Key to the larvae of synanthropic flies of Zaitsev, V. F. 1982. Microstructure of the labella of the fly Tadzhikistan. Keys for the fauna of the USSR. No. 28. proboscis. 1. Structure of the apparatus closing the Zoo!. Inst., Acad. Sci. USSR, Leningrad. pseudotracheae. - Ent. Rev., WasIl. 61(3):60-69. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae S9

9. Appendix (PI. 4C) ...... 4 Key to genera - CextendingtoMI (PI. lOA). Proboscis short or elongate...... 6 4. Lunulewithonepairofsetulae(asinPl.lB). 1. Vibrissa below or at the level oflower margin Palpus usually short and clavate, but some­ of eye; head in profile subquadrate; veliical times elongate. Basistemum very small and diameter of eye less than I.5x the horizontal shaped like a nanow triangle (PI. SF). Cross­ diameter (PI. ID). Frons in male as wide as in vein dm-cu absent (PI. 10C). Hind femur female. Wing with subcostal notch seldom without a strong seta anteroventrally ...... evident, never well-developed; R4 + 5 and ...... Paramyia Williston MI not or only slightly converging at tips; - Lunule bare. Palpus long and thin (PI. 4C). distal margin of anal cell (CuA2) usually Basistemum small and subquadrate (PI. SE, rounded (PI. lOG) (if not, then epistoma G). Crossvein dm-cu present or absent. Hind extended dorsally, subtriangular). Abdominal femur with a strong seta anteroventrally on tergites of male without silvery microto- distal half ...... 5 mentum ...... 2 5. Crossvein dm-cu absent. Arista slender, - Vibrissa well above lower margin of eye; nearly bare ...... Xenophyllomyza Ozerov head in profile rounded ventrally, distinctly - Crossvein dm-cu present. Arista thickened higher than long. Vertical diameter of eye at base and often densely long-haired ...... usually at least 1.5x the horizontal diameter ...... Aldrichiomyza Hendel (PI. IF). Frons in male nanower than in 6. Proboscis elongate; labella, when open, female. Wing with subcostal notch usually together at least 2x as long as broad, each evident (PI. lOJ), sometimes strongly so, labellum with 2 pseudotracheae (PI. 6D- with basal sector of C ending in an acute E) ...... 7 lappet; R 4 + 5 and M 1 sometimes converging - Proboscis short; labella, when open, together at tips; distal margin of anal cell meeting anal less than 2x as long as broad, each labellum vein in sharp angle (PI. 101). Abdominal with 4 pseudotracheae (PI. 7B) ...... S tergites of male often with silvery microto- 7. Only 2 lateroc1inate orbital setae (PI. 2C). mentum ...... 14. Milichiinae Basistemum reduced to a nanow triangle 2. Postocellar setae cruciate or so strongly (PI. SF). Hind femur without a strong seta converging that the tips nearly meet (PI. 1C). anteroventrally ...... Neophyllomyza Melander Basistemum small, without pre coxal bridge - Three lateroc1inate orbital setae (PI. IB). (PI. SE-G). First tarsomere of hind leg often Basistemum small and sub quadrate (PI. SE, I with a posterior comb of setae ventrally~g: G). Hind femur with a strong seta antero­ . ~ ...... 3. Phyllomyzmae ventrally on distal halL ... Stomosis Melander - Postocellar setae only slightly converging or S. Two postpronotal setae. Bases of antennae parallel (PI. 2E). Basistemum large and V­ widely separated, face with flatplate between shaped, sometimes with a pre coxal bridge antennae. Two or 3 frontal setae. Palpus en- (PI. SA-D). First tarsomere of hind leg with­ larged ...... Microsimus Aldrich out a posterior comb of setae ventrally ...... - One postpronotal seta. Antennae approxi­ ...... 11. Madizinae mated and not separated by a flat plate; 2 3. C extending to R4 + 5 or only slightly beyond frontal setae. Palpus enlarged or not ...... 9 (PI. 10C). Proboscis elongate; labella, when 9. First flagellomere in male as large as in open, together at least 2x as long as broad female. Mid femur with an anterodorsal 90 Irina Brake ENT. SCAND. SUPPL. S7 (2000)

ral organ slightly proximal of the middle (PI. l1A) ...... Madiza Fallen (PI. I1F-G) ...... Genus nov. 14. Male with all orbital and frontal setae re­ - First flagellomere in male usually larger than clinate; female with lower orbital seta latero­ in female. Mid femur without a femoral proclinate, middle orbital seta latero-rec1i- organ ...... 10 I nate, upper orbital seta~ absent, and frontal 10. Three lateroclinate orbital setae. Occiput not setae medioclinate. Wing oval (PI. lOG); strongly concave when viewed from above. subcostal break in costa not developed into a Palpus and first flagellomere in male usually notch. Abdominal tergites of male neve!' greatly enlarged, longer than broad. A pair of without silvery microtomentum ...... usually small setae on lunule ...... Enigmilichia Deeming ...... Phyllomyza Fallen - Male and female with lower orbital seta - Two or 3 lateroclinate orbital setae. Occiput usually latero-proc1inate, middle orbital seta strongly concave when viewed from above. f usually latero-reclinate, upper orbital set¥ Palpus not unusually enlarged; pedicel and medio-reclinate or absent, and frontal setae first flagellomere subequal in length, first medioclinate or absent (PI. 2A). Wing trian­ flagellomere broader than long, sub quadrate. gular (PI. 1OJ); subcostal break in costa deve­ A pair of strong setae on lunule ...... loped into a notch. Abdominal tergites of ...... Costa lima Sabrosky male often with silvery microtomentum .... IS 11. Frons with only one orbital seta (Pl. 2E) and IS. Frons with eight to ten pairs of uniformly without interfrontal setae or hairs ...... arranged frontal and orbital setae, all erect ...... Litometopa Sabrosky and reclinate, with no distinction between - Frons with 2 orbital setae (PI. 2B-D) and orbital and frontal setae (PI. IF). Proboscis with a paired row of interfrontal setae ...... 12 usually exceptionally long and slender, more 12. Frons with a paired row of interfrontal setae than 2x as long as head is high. Abdominal situated on grey sc1erotized interfrontal tergites of male without silvery microtomen- stripes; these stripes and concolorous frontal tum ...... Eusiphona Coquillett and orbital plates and ocellar triangle - Frons with orbital setae clearly distinguished demarcating a dull to velvet-black M-shaped from frontal setae, not unifom11y arranged. area (PI. 9A) ...... Desmometopa Loew Proboscis less than 2x as long as head is high, - Frons without an M-shaped area ...... 13 usually short (PI. SA). Abdominal tergites of 13. Lunule extended ventrally along mid-line of male often with silvery microtomenum ..... 16 face as a flat plate between the antennae, 16. Anepistemum with a row of strong setae almost reaching lower margin of face (PI. posteriorly. Three orbital setae and usually 3B); epistoma usually extended dorsally, three frontal setae, but sometimes 2 or up to subtriangular. Anepistemum sometimes with 6 frontal setae ...... Pholeomyia Bilimek hairs; anepimeron with 0-2 setae. Hind tibia - Anepistemum bare. Two or 3 orbital setae of male sometimes greatly broadened (PI. and no or 2 frontal setae ...... 17 lIE). Hind tibia in both sexes without tibial . 17. Posteriormarginofeyenotchedoremarginate organ...... Leptometopa Becker \ ~ ...... 18 - Lunule not extended towards lower margin - Posterior margin of eye entire (Milichia of face (PI. 3D); epistoma not extended Meigen) ...... 19 dorsally. Anepistemum and anepimeron bare. 18. Eyes more than I.Sxashighas wide. Labella, Hind tibia not enlarged in male. Hind tibia when open, together less than 2x as long as with tibial organ posteriorly in both sexes broad ...... Milichiella Giglio-Tos ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 91

- Eyes less than 1.5x as high as wide. Labella, confused with the true orbital and frontal when open, together more than 2x as long as setae. R4 + 5 and Ml parallel...... broad ...... Ulia Becker ...... Milichia speciosa-group 19. Usually no frontal setae in both sexes, and in - Female with 2 medioclinate frontal setae and some species 2 medioclinate frontal setae in 2 latero-reclinate or latero-proclinate orbital both sexes. Lower orbital seta in both sexes setae. Male with 1 lower mediociinate and 3 latero-proclinate, middle orbital seta latero­ upper medio-reciinate frontal setae and 1 reciinate, and upper orbital seta medio­ latero-reciinate orbital seta. R4 + 5 and Ml reciinate if present. In some species frontal converging at tips ...... and orbital hairs enlarged so that they can be ...... Milichia distinctipennis-group 92 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

10. Appendix ry(ies). zeus Author(s), year. Publication: page (genus Catalogue of world Milichiidae name in original description). Type-locali­ ty(ies): Town, province (country). (Anno­ This catalogue is based on the catalogue ofthe tation(s) such as "preocc. "). Diptera of America North of Mexico (Sabrosky 1965a), the catalogue ofthe Diptera oftheAmericas References given after each name are to the south of the United States (Sabrosky 1973), the original proposal of that name. Additional (secon­ catalogue of the Diptera of the Oriental Region dary) references are given in brackets when an (Sabrosky 1977), the catalogue of the Diptera of author described the same taxon as new in a different the Afrotropical Region (Sabrosky 1980), the work. The boundaries ofthe zoogeographic regions catalogue of the Diptera of the Australasian and correspond to the coverage of the catalogues Oceanian Regions (Sabrosky 1989), the catalogue mentioned above. Annotations added by me are in ofPalaearctic Diptera (Papp 1984a), and additional square brackets. papers dealing with Milichiidae (e. g. the checklist Suprageneric taxa (subfamilies) are ananged in of the Czech and Slovak Republics by Rohlicek the systematic order derived from the phylo genetic 1997, the checklist of Great Britain by Chandler tree of the Milichiidae (see the main pali of this 1998, the checklist of Switzerland by Papp 1998, paper). Within a taxon, the subordinate taxa are and the checklist of Israel by A. Freidberg, pers. listed alphabetically, i.e. genera within a subfamily, comm.). species within a genus. Synonyms are listed chro­ The format I have adopted for this catalogue nologically; variant spellings (termed "errors" follows that of the catalogue ofPalaearctic Diptera herein) and misidentifications are placed at the end (Papp 1984a). Details are illustrated in the following of the synonyms in alphabetical order. hypothetical examples ofgeneric and species entries In the index genera and species, which are not (genus Xus and species Xus albus and the synonyms valid (for example synonyms) or which do not genus Yus and species Yus zeus). All valid generic belong to Milichiidae, are indicated in italics. For and species names in the catalogue are indicated in example: bold face type. zeus (Author(s), year), (genus name in original description), present genus.

Genus Xus Author(s), year Abbreviations for zoogeographic regions: AFR - Afrotropical Region, AUS - Australasian/Ocea­ Author(s), year. Publication: page (annotation(s) nian Region, NEA - Nearctic Region, NEO - such as "n.n."). Neotropical Region, ORl- Oriental Region, PAL Type-species: Xus albus Author( s), year. Pub­ - Palaearctic Region lication: page (method oftype designation). Other abbreviations: acc. - according to, comb. Yus Author(s), year. Publication: page (annota­ - combination by, des. - designated by (subse­ tion(s) such as "preocc. "). Type-species: quently designated by), distr. - distribution, excl. Yus zeus Author(s), year. Publication: page - exclusive, I.c. -loco citato (at the place cited), (method of type designation). mono - by monotypy, n.n. -new name (replacement name), 11f. - near, orig. des. - by original albus Author(s), year. Publication: page (genus designation, preocc. - preoccupied, s. - south, name in original description). Type-locali­ subs. mono - by subsequent monotypy, widesp. - ty(ies): Town, province (country). - Distri­ widespread. bution: Zoogeographic region: Count- ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae

Family Milichiidae Type-species: Microsimus luteus Aldrich, Subfamily Phyllomyzinae 1926. l.c.: 3 (orig. des.). Aldrichiomyza Hendel, 1914 luteus Aldrich, 1926. Proc. U.S. natn. Mus. 69::1 (Microsimus). Type-locality: Isiamas (Boli­ Hendel, 1914a. Ent. Mitt. 3 :73 (n.n. forAldrichiella via). - Distr.: NEO: Bolivia. Hendel, 1911). Type-species: Aldrichiella agromyzina Neophyllomyza Melander, 1913 Hendel, 1911. Wien. ent. Ztg 30:37 (orig. des.). Melander, 1913a. Jl. N.Y. ent. Soc. 21:243. Aldrichiella Hendel, 1911. Wien. ent. Ztg 30:35 Type-species: Neophyllomyza quadricornis (preocc. Vaughan, 1903 (Coelenterata)). Melander, 1913a. l.c.: 243 (orig. des.). Type-species: Aldrichiella agromyzina Vichyia Villeneuve, 1920. Bull. Soc. ent. Fr. 1920: Hendel, 1911. l.c.: 37 (orig. des.). 69. Type-species: Vichyia acyglossa Ville­ neuve, 1920. l.c.: 70 (mon.). agromyzina (Hendel, 1911). Wien. ent. Ztg 30:37 (Aldrichiella). Type-locality: Brookings acyglossa (Villeneuve, 1920). Bull. Soc. ent. Fr. (South Dakota). - Distr.: NEA: South 1920:70 (Vichyia). Type-locality: Vichy Dakota to Ontario, s. to Kansas and North (France). - Distr. : PAL: widesp. in Europe, Carolina. former USSR (North European tenitory), elephas (Hendel, 1913). Supplta ent. 2:108 Mongolia. (A ldrichiella). Type-localities: Anping & quercus Duda, 1935a. Natuurh. Maandbl. 24: Taihupu (Taiwan). - Distr.: PAL: North 39 (Neophyllomyza). Type-localities: Ha­ Korea; ORI: Taiwan. belschwerdt (Poland) & Gliwice (Poland). flaviventris Iwasa, 1997. Jap. J. Ent. 65:826 anuda (Curran, 1936). Proc. Calif. Acad. Sci. (4) (Aldrichiomyza). Type-locality: Futakuchi­ 22:46 (Desmometopa). Type-locality: onsen, Akiv, Miyagi Pref. (Japan). - Distr.: Anuda Is. - Distr.: AUS: Solomon Is. PAL: Japan. (Anuda Is.). longirostris Hendel, 1931. Bull. Soc. ent. Egypte, approximatonervis (Lamb, 1914). Trans. Linn. (N.S.) 15(2):71 (Aldrichiomyza). Type­ Soc. Lond. (2, Zool.) 16:365 (Desmome­ localities: Wadi Beida & Wadi Edeib, S.E. topa). Type-locality: Mahe (Seychelles).­ Deseli (Egypt). - Distr. : PAL: Egypt, Israel; Distr.: AFR: Ethiopia, Seychelles. AFR: Sudan. inimica Seguy, 1938. Mission scient. de l'Omo (4) 1938: 3 64 (Neophyllomyza). Type-locality: Costalima Sabrosky, 1953 Marakwet, Elgeyo Escarpment (Kenya).­ Sabrosky, 1953. Ent. News 64:40. Distr.: AFR: Kenya. Type-species: Costalima myrmicola Sabrosky, leanderi (Hendel, 1924). Dt. ent. Z. 1924:406 1953.l.c.: 41 (orig. des.). (Phyllomyza, subgenus Neophyllomyza). Type-locality: Kremsmunster, Oberoster­ myrmicola Sabrosky, 1953. Ent. News 64:41 reich (Austria). - Distr.: P AL:Austria, (Costalima). Type-locality: Vicosa, Minas CzechRep., France, Great Britain, Hungary, Gerais (Brazil). - Distr.: NEO: Brazil. Romania, Slovakia, Switzerland. magnipalpis (Williston, 1896). Trans. R. ent. Soc. Microsimus Aldrich, 1926 Lond. 1896:443 (Phyllomyza). Type-loca­ Aldrich, 1926. Proc. U.S. natn. Mus. 69:2. lity: St. Vincent (St. Vincent and the Grena- 94 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

dines). - Distr.: NEO: West Indies (St. (Paramyia). Type-locality: Grenada. - Dis­ Vincent). tr.: NEO: West Indies (Grenada). magnipalpus (enor) Melander, 1913. Jl. N.Y. nitens (Loew, 1869). Berl. ent. Z. l3:45 (Phyllo­ ent. Soc. 21:243. myza). Type-locality: Pennsylvania. - nitens Melander, 19l3a. Jl. N.Y. ent. Soc. 21:244 Distr.: NEA: South Dakota to Quebec, s. to (Neophyllomyza). Type-locality: Avon Florida, also British Columbia, Albelia, (Idaho). -Distr.: NEA: Idaho. Arizona; NEO: widesp. s. to Chile and quadricornis Melander, 19l3a. Jl. N.Y. ent. Soc. Brazil. 21 :243 (Neophyllomyza). Type-localities: Idaho, Washington, New Mexico & Phyllomyza Fallen, 1810 Louisiana. - Distr.: NEA: Michigan to New Hampshire, s. to New Mexico and Florida, Fallen, 1810. Specim. ent. novam Dipt.:20. also Alaska, Idaho, Washington. Type-species: Phyllomyzasecuricornis Fallen, approximata (Malloch, 19l3a). Proc. U.S. 1823. Phytomyzides et Ochtidiae Sveciae: 8 natn. Mus. 46: 138 (Phyllomyza). Type­ (subs. mon.). locality: Washington (District of Colum­ Hendelimyza Duda, 1935a. Natuurh. Maandbl. bia). 24:25 (as subgenus ofPhyllomyza) (preocc. wulpi (Hendel, 1907). Wien. ent. Ztg 26:242 Frey, 1927 ()). Type-species: (Desmometopa; n.n. for Agromyza Neophyllomyza tetragona Hendel, 1924. minutissima Wulp, 1897) Type-locality: Dt. ent. Z. 1924:407 (orig. des.). New Guinea (probably = Papua New Guinea). - Distr.: AUS: Papua New Guinea. aelleni Papp, 1984b. Revue suisse Zool. 91 :242 minutissima (Wulp in Kertesz, 1897). (Phyllomyza). Type-locality: Grotte d'Istri­ Tenneszetr. Fliz.20:611 (Agromyza). Type­ pura, Hanguranketa (Sri Lanka). - Distr.: locality: New Guinea (probably = Papua ORI: Sri Lanka. New Guinea). (Preocc. Zetterstedt, 1860). beckeri Kramer, 1920. Zool. Jb. Syst. 43:331 (Phyllomyza). Type-locality: Sachsen (Ger­ Paramyia Williston, 1897 many). - Distr.: PAL: Czech Rep., Ger­ many, Great Britain. Williston, 1897. Kans. Univ. Q. (A) 6:l. cavernae de Meijere, 1914. Tijdschr. Ent. 57:253 Type-species: Paramyia nigra Williston, 1897. (Phyllomyza). Type-locality: Gunung l.c.: 2 (mon.). Sewoe, Djocja (Java). - Distr.: OR!: Indonesia (Java), Malaysia. fumipennis Malloch, 1934. Dipt. ofPatag. and S. dilatata Malloch, 1914. AnnIs hist.-nat. Mus. natn. Chile 6 (5):465 (Paramyia). Type-locality: hung. 12:311 (Phyllomyza). Type-locality: Iquitos (Peru). - Distr.: NEO: Peru. Toyenmongai (Taiwan). - Distr.: ORI: hungarica Papp, 1993. AnnIs hist.-nat. Mus. natn. Taiwan. hung. 85:135 (Paramyia). Type-locality: donisthorpei Schmitz, 1923. Konowia 2:47 Aggtelek, Medveskeli (Hungary). - Distr.: (Phyllomyza). Type-localities: Darenth PAL: Hungary. Wood & Weybridge, Suney (England). - inconspicua de Meijere, 1916. Tijdschr. Ent. Distr.: PAL: Czech Rep., France, Great 59:211 (Paramyia). Type-locality: Tjibodas Britain, Hungary, Slovakia, Switzerland. (Java). - Distr.: ORI: Indonesia (Java). epitacta Hendel, 1914b. Supplta ent. 3 :97 (Phyllo­ nigra Williston, 1897. Kans. Univ. Q. (A) 6:2 myza). Type-localities: Chipun & Paroe ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae

(Taiwan). - Distr.: ORI: Taiwan. Sittard (Netherlands). - Distr.: PAL: Czech equitans (Hendel, 1919). Ent. Mitt. 8: 198 (Neo­ Rep., Great Britain, Hungary, Netherlands, phyllomyza). Type-localities: Bisambergm. Poland, Romania, Slovakia. Vienna (Austria) & Freiheitsau nr. Troppau lucens Hendel, 1924. Dt. ent. Z. 1924:40S (Czech Rep.). - Distr.: PAL: Austria, Czech (Phyllomyza). Type-locality: Rapall() Rep., Finland, Germany, Great Britain, (Italy). - Distr. : PAL: Italy. Hungary, Norway, Slovakia, Switzerland; lutea de Meijere, 1914. Tijdschr. Ent. 57:25) fonner USSR: North and Central European (Phyllomyza). Type-locality: Salatiga territories, ?Far East. (Java). - Distr.: ORI: Indonesia (Java). fagicola Hendel, 1924. Dt. ent. Z. 1924:408 luteipalpis Malloch, 1914. AnnIs hist.-nat. Mus. (Phyllomyza). Type-locality: Waldviertel, natn. hung. 12:310 (Phyllomyza). Type Niederosterreich (Austria). locality: Takao (Taiwan). - Distr.: ORI: lasiae(Duda, 1935a).Natuurh. Maandbl. 24:37 Taiwan. (Hendelimyza). Type-locality: not given melania (Hendel, 1919). Ent. Mitt. 8:198 (Nen (England). phyllomyza). Type-locality: Unterberg, flavipalpis de Meijere, 1914. Tijdschr. Ent. 57 :252 Niederosterreich (Austria). - Distr.: P /\ I,: (Phyllomyza). Type-localities: Salatiga, Austria, Czech Rep., Hungary, Slovakia, Semarang, Tandjong Priok & Wonosobo Switzerland. (Java). - Distr.: ORI: Indonesia (Java); milnei Steyskal, 1942. Ent. News 53:84 (Phyl/II AUS: ?Solomon Is. myza). Type-locality: Mountain Lake flavitarsis (Meigen, 1830). Syst. Beschr. 6: 172 (Virginia). - Distr.: NEA: Illinois, Mary (Agromyza). Type-locality: not given land, North Carolina, Virginia. (?Aachen, Germany). - Distr.: PAL: mongolica Papp, 1976. Acta zool. hung. 22:37() Belgium, Czech Rep., Germany, Great (Phyllomyza). Type-locality: "Bulgall Britain, Hungary, Slovakia, Sweden, aimak:30 km NNW von Somon Daschill Switzerland. cilen" (Mongolia).-Distr.: PAL: Mongolia. tarsata (Zetterstedt, 1848). Dipt. Scand. 7:2723 nigripalpis de Meijere, 1914. Tijdschr. Ent. 57:2S.1 (Milichia). Type-localities: Oland & "Ofa" (Phyllomyza). Type-locality: Salaliga (Sweden). (Java). - Distr.: ORI: Indonesia (Java). formicae Schmitz, 1923. Konowia 2:46 nudipalpis Malloch, 1914. AnnIs hist.-nat. MilS. (Phyllomyza). Type-locality: Weybridge, natn. hung. 12:311 (Phyllomyza). Type Surrey (England). - Distr.: PAL: Denmark, locality: Takao (Taiwan). - Distr.: () IU: France, Great Britain. Taiwan. fuscogrisea (Seguy, 1933). Mems Estud. Mus. pallida de Meijere, 1940. Ent. Ber., Arnst. 10 (No, zool. Univ. Coimbra 67:50 (Neophyllo­ 232):222 (Phyllomyza). Type-Iocalily: myza). Type-locality: N ova-Choupanga, m. Nunspeet (Netherlands). - Distr.: 1'/\1: Chemba (Mozambique). - Distr.: AFR: Netherlands. (A doubtful species aec. "lIpP Mozambique. 1984.) hirtipalpis Malloch, 1913a. Proc. U.S. natn. Mus. rubricornis Schmitz, 1923. Konowia ):,1>1 46: 137 (phyllomyza). Type-locality: Plum­ (Phyllomyza). Type-locality: BlidllPl'SI mers Is. (Maryland). - Distr.: NEA: Mary­ (Hungary).-Distr.: PAL: Czech Rep., ("l'lIl land, Tennessee, Texas, Virginia. Britain, Hungary, Switzerland, /'OIllll'1 longipalpis (Schmitz, 1924). Natuurh. Maandbl. USSR: North European telTitory. 13: 16 (Neophyllomyza). Type-locality: securicornis Fallen, 1823. PhytoI11Y!',idcs l'I 96 Irina Brake ENT. SCAND. SUPPL. 57 (2000) t Ochti~if:lae Sveciae:8 (Phyllomyza). Type­ Melbourne (Victoria). - Distr.: AUS: Vic­ locality: Esperod (Sweden). - Distr.: PAL: toria. widesp. in Europe; former USSR: North innominata (Williston, 1896). Trans. R. ent. Soc. and Central European tenitories (Russian Lond.1896:443 (Agromyza). Type-locality: SFSR), Far East; NEA: Wisconsin to St. Vincent (St. Vincent and the Grena­ Quebec and Maine, s. to North Carolina, dines). - Distr.: NEO: Brazil, Colombia, also British Columbia and Alberta. Costa Rica, El Salvador, Honduras, Mexico, jlavitarsis (Meigen, 1830). Syst. BeschI'. 6: 107 Panama and Canal Zone, West Indies (St. (Opomyza). Type-locality: not given Vincent, Trinidad); NEA: Arizona. (?Aachen, Germany). luteola (Coquillett, 1902). n. N.Y. ent. Soc. jlavipes (Meigen, 1830). Syst. Beschr. 6: 108 10:188 (Desmometopa). Type-locality: (Opomyza). Type-locality: not given Williams (Arizona). (?Hamburg district, Gelmany). rufula (Frey, 1918). Gfvers. finska VetenskSoc. silesiaca (Duda, 1935a). Natuurh. Maandbl. 24:38 Forh. 60 (1917-1918, Afd. A, no. 14):18 (Hendelimyza). Type-locality: Habel­ (Siphonomyiella). Type-locality: Rio de schwerdt (Poland). - Distr. : PAL: Poland; Janeiro (Brazil). - Distr.: NEO: Brazil. fonner USSR: North European tenitory. vittata Malloch, 1925. Proc. Linn. Soc. N.S.W. tenebrosa Bmnetti, 1924. Rec. Indian Mus. 26: 104 50:89 (Stomosis). Type-locality: Sydney (Phyllomyza). Type-locality: Siju Cave, (New South Wales). - Distr.: AUS: Garo Hills, Assam (India). - Distr.: OR!: Australian Capital Tenitory, New South India. Wales. tetragona Hendel, 1924. Dt. ent. Z. 1924:407 (Phyllomyza). Type-localities: Vienna Xenophyllomyza Ozerov, 1992 (Austria) & Legnica (Poland). - Distr.: PAL: Austria, Gennany, Hungary, Poland, Ozerov, 1992. Vestnik zoo 1. 1992(2):18. Slovakia, Switzerland. Type-species: Xenophyllomyza deserticola Ozerov, 1992. l.c.: 19 (orig. des.). Stomosis Melander, 1913 deserticola Ozerov, 1992. Vestnikzool. 1992(2): 19 Melander, 1913a. Jl. N.Y. ent. Soc. 21:242. (Xenophyllomyza). Type-locality: Batchiz Type-species: Desmometopa luteola Coquil­ (Turkmenistan). - Distr.: PAL: Turk­ lett, 1902. Jl. N.Y. ent. Soc. 10:188 (orig. menistan. des.) (= innominata (Williston, 1896)). SiphonomyiellaFrey, 1918. Gfvers. finska Vetensk­ Subfamily Madizinae Soc. Forh. 60 (1917-1918, Afd. A, no. Desmometopa Loew, 1866 14): 16. Type-species: Siphononzyiella rufula Frey, 1918.l.c.: 18 (orig. des.). Loew, 1866. Bed. ent. Z. 9 (1865) (Centuria 6, No. 96): 184. flava Sabrosky, 1958a. Ent. News 69:172 Type-species: Agromyza m-atrum Meigen, (Stomosis). Type-locality: Lafayette (Indi­ 1830. Syst. Beschr. 6:170 (des. Hendel, ana). -Distr.: NEA: Michigan to Virginia, 1903. Wien. ent. Ztg 22:251) (= sordida s. to Alabama and Florida. (Fallen, 1820)). flavoscutellata Malloch, 1925. Proc. Linn. Soc. Platophrymia Williston, 1896. Trans. R. ent. Soc. N.S.W. 50:88 (Stomosis). Type-locality: Lond. 1896:426. Type-species: Platophry- ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 97

mia nigra Williston, 1896.l.c.: 426. (mon.) flavicoxa Hendel, 1932. Konowia II: 14:1 (= tarsalis Loew, 1866) (Desmometopa). Type-locality: Mis. Ta Liodesma Duda, 1935a. Natuurh. Maandbl. 24:25 caagle (Argentina). - Distr.: NEO: Argcn (as subgenus of Desmometopa). Type­ tina, Paraguay, Uruguay. species: Liodesma atra Duda, 1935a. l.c.: flavipalpis Sabrosky, 1983. Contr. Am. ent. Ins!. 25 (orig. des.) (=sordida (Fallen, 1820)). 19(8):49 (Desmometopa). Type-locality: Liodesll1ometopa Duda, 193 5a. N atuurh. Maandbl. Jaluit Atoll, Majurirek Is. (Marshall Is.). 24:24 (invalid: first published as a synonym Distr.: AUS: Caroline Is., Indonesia (Irian of Liodesma Duda). Jaya), Marshall Is., Belau, Vanuatu. floridensis Sabrosky, 1983. Contr. Am. ent. Ins!. aczeli Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):33 (Desmometopa). Type-locality: 19(8): 62 (Desmometopa). Type-locality: Sanford, Seminole Co. (Florida). - Distr.: Vista Flores, Mendoza (Argentina). - Distr.: NEA: Florida, Georgia. NEO: Argentina. glaucanota Sabrosky, 1983. Contr. Am. ent. Inst. aldabrae Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):32 (Desmometopa). Type-locality: 19(8):53 (Desmometopa). Type-locality: Corozal Town (Belize). - Distr.: NEO: South Is. (AldabraISeychelles). - Distr.: Belize, Mexico, Panama. AFR: Seychelles. gressitti Sabrosky, 1983. Contr. Am. ent. Inst. argentinica Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):54 (Desmometopa). Type-locality: 19(8):59 (Desmometopa). Type-locality: JaluitAtoll, Jabor Is. (Marshall Is.). - Distr.: Salta, Urundel (Argentina). - Distr.: NEO: AUS: Caroline Is., Gilbert Is., Marshall Is., Argentina, Peru. Solomon Is., Vanuatu. atypica Sabrosky, 1983. Contr. Am. ent. Inst. inaurata Lamb, 1914. Trans. Linn. Soc. Lond. (2, 19(8):39 (Desmometopa). Type-locality: Zool.) 16:363 (Desmometopa). Type­ Las Cumbres, Panama Province (Panama). localities: Seychelles. - Distr.: widesp. in - Distr.: NEO: Ecuador, Panama, Peru. AFR and AUS; NEO: Brazil, El Salvador, blantoni Sabrosky, 1983. Contr. Am. ent. Inst. Guyana, WestIndies (Bahamas, Dominica, 19(8):35 (Desmometopa). Type-locality: Ft. Puerto Rico, Virgin Is.). Kobbe, Camal'on, Canal Zone (Panama).­ semiaurata Sabrosky, 1958b. Stuttg. Beitr. Distr.: NEO: Costa Rica, EI Salvador, Naturk. 4:4 (Desmometopa). Type-locality: Mexico, Panama. Msingi (Tanzania). ciliata Hendel, 1919. Ent. Mitt. 8:200 (Desmome­ ciliata (misidentification) Malloch, 1924. Proc. topa). Type-locality: Sydney (New South Linn. Soc. N.S.W. 49:336. Wales). - Distr.: AUS: Australian Capital indistincta Sabrosky, 1983. Contr. Am. ent. Inst. TelTitory, New South Wales, Queensland, 19(8):34 (Desmometopa). Type-locality: South Australia, Western Australia. Iquitos (Peru). - Distr.: NEO: Colombia, discipalpis Papp, 1993. AnnIs hist.-nat. Mus. natn. Costa Rica, Ecuador, Peru. hung. 85:133 (Desmometopa). Type-loca­ interfrontalis Sabrosky, 1965b. Stuttg. Beitr. lity: Tihany, Csucshegy (Hungary). - Distr.: Naturk. 138:3 (Desmometopa). Type-loca­ PAL: Hungary. lity: Makoa (Tanzania). - Distr.: AFR: evanescens Sabrosky, 1983. Contr. Am. ent. Inst. Cameroun, Ivory Coast, Liberia, Namibia, 19(8):28 (Desmometopa). Type-locality: Nigeria, Oman, Tanzania, Uganda, Zaire. Las Cumbres, Panama Province (Panama). kandyensis Sabrosky, 1983. Contr. Am. ent. Inst. - Distr.: NEO: Panama. 19(8): 54 (DesnlOmetopa). Type-locality: 98 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Udawattakele, Kandy District (Sri Lanka). Joshua Tree National Monument (Cali­ - Distr.: ORI: Sri Lanka. fornia). - Distr.: NEA: Arizona, California, Iatigena Sabrosky, 1983. Contr. Am. ent. Inst. District of Columbia, Georgia, Iowa, Kan­ 19(8):24 (Desmometopa). Type-locality: sas, Mexico, New York, Texas, Utah. Big Bend National Park (Texas). - Distr.: nigeriae Sabrosky, 1983. Contr. Am. ent. Inst. NEA: California, Texas. 19(8) :21 (Desmometopa). Type-locality: Ieptometopoides Sabrosky, 1983. Contr. Am. ent. Olokemeji (Nigeria). - Distr.: AFR: Nigeria. Inst. 19(8):40 (Desmometopa). Type­ nigrohalteralis Sabrosky, 1983. Contr. Am. ent. locality: Suakoko (Liberia). -Distr.: AFR: Inst. 19(8):31 (Desmometopa). Type-loca­ Camerouon, Ghana, Liberia, Nigeria, lity: Villa Padre Monti, Tucuman-Burru­ ?South Africa; PAL: Tunisia. yacu (Argentina). - Distr. : NEO: Argentina. Iucidifrons Sabrosky, 1983. Contr. Am. ent. Inst. nudigena Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):23 (Desmometopa). Type-locality: 19(8):48 (Desmometopa). Type-locality: Simla (Trinidad and Tobago). - Distr.: Bakau (Gambia). - Distr.: AFR: Gambia, NEO: WestIndies (Trinidad). Ivory Coast, Nigeria, Sierra Leone. magnicornis Sabrosky, 1983. Contr. Am. ent. obscurifrons Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):51 (Desmometopa). Type-loca­ Inst. 19(8):37 (Desmometopa). Type-loca­ lity: Ibadan (Nigeria). - Distr.: AFR: Ivory lity: David, Chiriqui (Panama). - Distr.: Coast, Nigeria. NEO: Colombia, Costa Rica, Ecuador, El melanderi Sabrosky, 1983. Contr. Am. ent. Inst. Salvador, Mexico, Panama, West Indies 19(8):60 (Desmometopa). Type-locality: (Tobago). Verdemont, San Bernardino Co. (Califor­ parafacialis Sabrosky, 1983. Contr. Am. ent. Inst. nia). - Distr.: NEA: Arizona, California, 19(8):38 (Desmometopa). Type-locality: Mexico, Texas, Utah. Austin (Texas). - Distr.: NEA: Mexico, meridionalis Sabrosky, 1983. Contr. Am. ent. Texas. Inst. 19(8):34 (Desmometopa). Type-loca­ philippinensis S abro sky, 1983. Contr. Am. ent. lity: Nova TeutOnia, Santa Catarina (Brazil). Inst. 19(8):52 (Desmometopa). Type­ - Distr.: NEO: Argentina, Bolivia, Brazil, locality: Manila (Philippines). - Distr.: ORr: Peru, Uruguay. Philippines. microps Lamb, 1914. Trans. Linn. Soc. Lond. (2, pleuralis Sabrosky, 1983. Contr. Am. ent. Inst. Zool.) 16:364 (Desmometopa). Type-loca­ 19(8):21 (Desmometopa). Type-locality: lity: Seychelles. - Distr.: widesp. in ORr Bakau (Gambia). - Distr.: AFR: Gambia, and bordering areas of PAL (Afghanistan, Nigeria. Nepal, China, Japan); AFR: Cameroun, postorbitalis Sabrosky, 1983. Contr. Am. ent. Seychelles, Tanzania. Inst. 19(8):26 (Desmometopa). Type-loca­ m-nigrum (Zetterstedt, 1848). Dipt. Scand. 7:2743 lity: Mabungo, Kigezi Province (Uganda). (Agromyza). Type-localities: Vadstcna & - Distr.: AFR: Kenya, Uganda. "Liiderbro" (Sweden). - Distr.: cosmopoli­ propeciliata Sabrosky, 1983. Contr. Am. ent. Inst. tan, excl. oceanic islands. 19(8): 51 (Desmometopa). Type-locality: nUotica Becker, 1903. Mitt. zool. Mus. Berl. Pahang (Malaysia). - Distr.: ORr: Indonesia 2: 188 (Desmometopa). Type-locality: "Nile (Java), Malaysia, Thailand. Valley" (Egypt). saguaro Sabrosky, 1983. Contr. Am. ent. Inst. nearctica Sabrosky, 1983. Contr. Am. ent. Inst. 19(8):55 (Desmometopa). Type-locality: 19(8):58 (Desmometopa). Type-locality: Saguaro National Monument, Pima Co. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 99

(Arizona). - Distr.: NEA: Arizona, Cali­ terminalis Sabrosky, 1983. Contr. Am. ent. Inst. fornia. 19(8):36 (Desmometopa). Type-locality: singaporensis Kertesz, 1899. Tenneszetr. Fiiz. Korol' Island (Belau). - Distr.: AUS: Belau. 22: 194 (Desmometopa). Type-locality: varipalpis Malloch, 1927. Proc. Linn. Soc. N. S. W. Singapore. - Distr.: widesp. in AFR, AUS, 52:7 (Desmometopa). Type-locality: and ORI; PAL: Afghanistan; NEO: Brazil, Bourke (New South Wales). -Distr.: PAL: West Indies (Puerto Rico). Afghanistan, Egypt, Germany, Great palpalis de Meijere, 1914. Tijdschr. Ent. Britain, Spain; widesp. in AFR, A US, NEA, 57:251 (Desmometopa). Type-localities: NEO, and OR!. Salatiga (Java) & Deli (Sumatra). singaporensis (misidentifications) authors, not tristicula Hendel, 1914b. Supplta ent. 3:96 Kertesz (see Sabrosky 1983). (Desmometopa). Type-locality: Anping tarsalis (misidentifications) authors, not Loew (Taiwan). (see Sabrosky 1983). m-nigrul11 (misidentification) Illingworth woldai Sabrosky, 1983. Contr. Am. ent. Inst.19(8): 1926. Proc. Hawaii. ent. Soc. 6:224. 29 (Desmometopa). Type-locality: Las tarsalis (misidentifications) authors, not Loew Cumbres, Panama Province (Panama). - (see Sabrosky 1983). Distr.: NEO: Panama. sordida (Fallen, 1820). Oscinides Sveciae: 10 (Madiza). Type-locality: "Esperod" Leptometopa Becker, 1903 (Sweden). - Distr.: widesp. in PAL and NEA. Becker, 1903. Mitt. zool. Mus. Berl. 2(3):188. m-atrum (Meigen, 1830). Syst. Beschr. 6: 170 Type-species: Leptometopa rufiFons Becker, (Agromyza). Type-locality: not given 1903. l.c.: 188 (mon.). (?Aachen, Germany). Macrurus Lioy, 1864. Atti Ist. veneto Sci. (3) atra (Duda, 1935a). Natuurh. Maandbl. 24:38 9: 1313 (preocc. Bloch, 1780 (Pisces)). (Liodesma). Type-locality: Habelschwerdt Type-species: Agromyza latipes Meigen, (Poland) & Sankt Wendel (Germany). 1830. Syst. BeschI'. 6: 177 (mon.). srilankae Sabrosky, 1983. Contr. Am. ent. Inst. Hypaspistomyia Hendel, 1907. Wien. ent. Ztg 19(8):50 (Desmometopa). Type-locality: 26:240. Type-species: Hypaspistomyia Udawattakele, Kandy District (Sri Lanka). coquilletti Hendel, 1907. l.c.: 240 (mon.). - Distr.: ORI: Sri Lanka, ?Philippines. Paramadiza Malloch, 1913a. Proc. U. S. natn. Mus. stilbopleura Sabrosky, 1983. Contr. Am. ent. Inst. 46:136 (preocc. Melander, 1913). Type­ 19(8):59 (Desmometopa). Type-locality: species: Desmometopa halteralis Coquil­ Sao Paulo, Nova Teutonia, Santa Catarina lett, 1900. Proc. U.S. natn. Mus. 22:276 (Brazil). - Distr.: NEO: Brazil. (orig. des.). tarsalis Loew, 1866. Berl. ent. Z. 9 (1865) (Centuria Mallochiella Melander, 1913b. Psyche, Camb. 6, No. 96):184 (Desmometopa). Type­ 20: 169 (n.n. for Paramadiza Malloch, locality: Cuba. - Distr.: AUS: widesp. on 1913). Type-species: Desmometopa oceanic islands; NEA: Arizona, Texas; halteralis Coquillett, 1900. Proc. U.S. natn. NEO: widesp. in northern part. Mus. 22:276 (automatic). nigra (Williston, 1896). Trans. R. ent. Soc. Prodesmometopa Hendel, 1914b. Supplta ent. 3: 97. Lond. 1896:426 (Platophlymyia). Type­ Type-species: Agromyza latipes Meigen, locality: St. Vincent (St. Vincent and the 1830. Syst. Beschr. 6:177 (orig. des.). Grenadines) . Cnemoplegas Enderlein, 1927. Stettin. ent. Ztg 100 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

88: 108. Type-species: Agromyza latipes widesp. Meigen, 1830. Syst. Beschr. 6: 177 (orig. kaszabi Papp, 1976. Acta zool. hung. 22:371 (Lep­ des.). tometopa). Type-locality: "Sudgobi aim­ Desmometopina Cunan, 1930. Bull. Am. Mus. ak: SW Rand des Salzsees Dund gol (am nat. Hist. (1931) 61 :81. Type-species: Agro­ "alten" Somon Gurban-tes)" (Mongolia).­ myza latipes Meigen, 1830. Syst. Beschr. Distr. : PAL: Mongolia; AFR: Kuwait, Sau­ 6:177 (orig. des.). di Arabia. Dicraeoptera Duda, 1932. 61. Chloropidae,Fliegen lacteipennis (Hendel, 1913). Supplta Ent. 2:108 pal. Reg. 6(1 ):34. Type-species: Siphonella (Madiza). Type-localities: Anping & Tainan niveipennis Strobl, 1900a. Wiss. Mitt. Bosn. (Taiwan). - Distr.: ORI: Taiwan. Herzeg. 7:631 (orig. des.). latipes (Meigen, 1830). Syst. Beschr. 6: 177 Rhynchomadiza Enderlein, 1934. Sber. Ges. naturf. (Agromyza). Type-localities: Berlin & Freunde Berl. 1933 :427. Type-species: Solingen (Germany). - Distr.: widesp. in Rhynchomadizajlaviceps Enderlein, 1934. AFR, NEA, NEO, ORI, and PAL. I.c.: 427 (orig. des.). annulimana (von Roser, 1840). ConespBI. wlirtt. landw. Ver. Stuttg. (N.S.) 1(1):63 aelleni Papp, 1978. Revue suisse Zool. 85:104 (Agromyza). Type-locality: not given (Leptometopa). Type-locality: Yate (New (Wiilitemberg, Germany). Caledonia). - Distr.: AUS: New Caledonia. annulitarsis (von Roser, 1840). ConespBI. albipennis (Lamb, 1914). Trans. Linn. Soc. Lond. wlirtt. landw. Ver. Stuttg. (N.S.) 1(1):63 (2, Zool.) 16:366 (Madiza). Type-locality: (Agromyza). Type-locality: not given Mahe (Seychelles). - Distr.: AFR: (Wurttemberg, Gelmany). Seychelles. annulitarsis (Zetterstedt, 1848). Dipt. Scand. beardsleyi Hardy & Delfinado, 1980. Insects of 7:2674 (Madiza). Type-localities: "Lundae Hawaii 13:358 (Leptometopa). Type-loca­ ... e Dania". lity: Lisianski Is. (Hawaiian Is). - Distr.: orillia (Curran, 1927). Can. Ent. 59:50 AUS: Hawaiian Is., Pitcairn Is. (Mallochiella). Type-locality: Ontario. broersei de Meijere, 1946. Tijdschr. Ent. mallochi Sabrosky, 1989. Cat. Dipt. Australasia & 87 (1944): 17 (Leptometopa). Type-locality: Oceania:556 (Leptometopa, n.n. for Amsterdam (Netherlands). - Distr.: PAL: albipennis (Malloch, 1924)). Type-locality: Netherlands. (a doubtful species acc. Papp New South Wales (Australia). - Distr.: 1984). AUS: widesp. coquilletti (Hendel, 1907). Wien. ent. Ztg 26:241 albipennis Malloch, 1924. Proc. Linn. Soc. (Hypaspistomyia). Type-locality: Aden N.S.W. 49:336 (Hypaspistomyia). Type­ (Yemen). - Distr.: AFR: Arabia, Nigeria, locality: New South Wales. (Preocc. Lamb, Sudan, Yemen. 1914). flaviceps (Enderlein, 1934). Sber. Ges. naturf. matilei Sabrosky, 1979. Mem. Mus. natn. Hist. FreundeBerl. 1933:427 (Rhynchomadiza). Type-locality: Alt-Hensis (Namibia). - Distr.: AFR: Namibia. halteralis (Coquillett, 1900). Proc. U.S. natn. Mus. 22:267 (Desmometopa). Type-locality: Utuado (Puerto Rico). - Distr.: NEO: Mexico, West Indies (Puerto Rico); NEA: ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 101 nilssoni Sabrosky, 1987. Proc. ent. Soc. Wash. Type-species: Litometopa glabriji'ons Sabros­ 89:242 (Leptometopa). Type-locality: Man­ ky, 1965b. 1.c.: 4 (orig. des.). draka (Madagascar). - Distr.: AFR: Benin, Cape Verde Is., Madagascar, ?Tanzania, glabrifrons Sabrosky, 1965b. Stuttg. Beitr. Naturk. ?Uganda, Yemen. 138: 4 (Litometopa). Type-locality: Maran­ niveipennis (Strobl,r91-l0a1. Wlss. Mitt. Bosn. gu (Tanzania). - Distr.: AFR: Rwanda, Herzegl.?r:l.lf(Siphonella). Type-localities: Tanzania. Dalmatia (Mostar, Spalato, Sebenico) (former Yugoslavia) & Triest (Italy). - Distr.: PAL: widesp.; NEO: West Indies Madiza Fallen, 1810 (Cuba). simplicipes (Becker, 1907a). Wien. ent. Ztg Fallen, 1810. Specim. ent. novam Dipt.: 19. 26:4 (Desmometopa). Type-locality: Hil­ Type-species: Madiza glabra Fallen, 1820. versum (Netherlands). Oscinides Sveciae: 9 (des. Hendel 1903. niveipennis fascifrons (Becker, 1907b). AnnIs Wien. ent. Ztg 22:251). hist.-nat. Mus. natn. hung. 5:548 (1910:664) Cleptoneria Lioy, 1864. Atti 1st. veneto Sci. (3) 9: (as Desmometopa fascifrons). Type­ 1120. Type-species: Madizaglabra Fallen, locality: Ajaccio, Corsica (France). - Distr.: 1820. Oscinides Sveciae: 9 (mon.). PAL: France. Polphorilla Enderlein, 1921. Zoo1. Anz. 52(8/9): pacifica Papp, 1984b. Revue suisse Zoo1. 91:243 231. Type-species: Piophila jlavitarsis (Leptometopa). Type-locality: Waituloa Meigen, 1830. Syst. Beschr. 6:383 (orig. Cave, Viti Levu (Fiji). - Distr.: AUS: Fiji. des.) (= glabra Fallen, 1820). pecki Papp, 1984b. Revue suisse Zoo 1. 91:245 Desmomyza CUlTan, 1934. Fam. Gen. N. Am. (Leptometopa). Type-locality: Waituloa Dipt.: 338. Type-species: Desmomyza COll­ Cave, Viti Levu (Fiji). - Distr.:AUS: Fiji. fusa Curran, 1934. 1.c.: 338 (orig. des.) (= rufifrons Becker, 1903. Mitt. zoo1. Mus. Ber1. 2: :slabra Fallen, 1820). 188 (Leptometopa). Type-locality: Suez (Egypt). - Distr.: PAL: Afghanistan, britannica Hennig, 1937. 60a. Milichiidae et Algeria, ?China (Tibet), Egypt, Iran, Israel, Camidae, Fliegen pal. Reg. 6(1):53 (Madi­ Libya, Romania, Spain (Canary Is.), za). Type-locality: Snailwell, Cambridge­ Tunisia, Turkey, former Yugoslavia; former shire (England). - Distr. : PAL: Great USSR: Soviet Middle Asia (Usbek SSR); Britain, Slovakia, Switzerland. AFR: Namibia, Oman, South Africa, eximia Papp, 1993. AnnIs hist.-nat. Mus. natn. Yemen. hung. 85:134 (Madiza). Type-locality: jlavipalpis Seguy, 1932. Annali Mus. civ. Bukk National Park, Szentlelek hg. Stor. nat. Giacomo Doria 55:502 (Lepto­ (Hungary). - Distr.: PAL: Hungary, metopa). Type-localities: Augila & Sebcha Slovakia. di Buema (Libya). glabra Fallen, 1820. Oscinides Sveciae: 9 (Madi­ latigenis (Hendel, 1933a). Dt. ent. Z. 1933:54 za). Type-locality: "Scania" (Ski'me), Viis­ (Hypaspistomyia). Type-localities: Arbe & tergotland (Sweden). - Distr.: PAL: Dalmatia (former Yugoslavia). widesp.; NEA: Washington. Litometopa Sabrosky, 1965 con/usa (Curran, 1934). Fam. Gen. N. Am. Dipt.: 338 (Desmomyza). Type-locality: Sabrosky, 1965b. Stuttg. Beitr. Naturk. 138:4. New York. 102 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

flavitarsis (Meigen, 1830). Syst. Beschr. 6:383 (Eusiphona). Type-locality: Canada de (Piophila). Type-locality: not given Gomez, Santa Fe (Argentina). - Distr.: (?Aachen, Germany). NEO: Argentina. rufitarsis (Meigen, 1838). Syst. Beschr. 7:384 (Gymnopa). Type-locality: not given Milichia Meigen, 1830 (?Aachen, Germany). pachymera Becker, 1908b. AnnIs hist.-nat. Mus. Meigen, 1830. Syst. Beschr. 6: 131. natn. hung. 6:320 (Madiza). Type-locality: Type-species: Milichiaspeciosa Meigen, 1830. Ugod (Hungary). - Distr.: PAL: Czech I.c.: 132 (des. Westwood 1840. Synopsis: Rep., Great Britain, Hungary, Israel, Slo­ 151). vakia, former Yugoslavia. Argyritis Latreille in Cuvier, 1829. La regne anima!..., ed. 2, 5: 518 (without included Subfamily Milichiinae species) (preocc . Hubner, 1816 (Lepido­ Enigmilichia Deeming, 1981 ptera)). Lobioptera Wahlberg, 1847. Ofvel's. K. Vetensk­ Deeming, 1981. Zool. J. Linn. Soc. 71:160. Akad. Forh. 4(9):259. Type-species: Type-species: Enigmilichia dimorphica Lobioptera ludens Wahlberg, 1847. I.c.: Deeming, 1981. l.c.: 162 (orig. des.). 259 (mon.). Prosaetomilichia de Meijere, 1909. Tijdschr. Ent. dimorphic a Deeming, 1981. Zool. J. Linn. Soc. 52: 170. Type-species: Prosaetomilichia 71: 162 (Enigmilichia). Type-locality: myrmecophila de Meijere, 1909.I.c.: 171 Mokwa, Niger State (Nigeria). - Distr.: (des. Sabrosky 1977. Cat. Dipt. Oriental AFR: Nigeria. Reg.: 274).

Eusiphona Coquillett, 1897 aethiops Malloch, 1913a. Proc. U.S. natn. Mus. 46: 133 (Milichia). Type-locality: Harlingen Coquillett, 1897. U.S. Dep. Agric., Div. Ent., Tech. (Texas). - Distr.: NEA: Mexico, Texas. Ser. 7:49. albomaculata (Strobl, 1900b). Wien. ent. Ztg 19:9 Type-species: Eusiphona mira Coquillett, (Lobioptera). Type-locality: Iativa(Spain). 1897.l.c.: 49 (orig. des.). - Distr.: PAL: ?Israel, Marocco, Spain; AFR: Yemen. cooperi Sabrosky, 1955. Ent. News 66:170 (Eusi­ angustifrons Bezzi, 1928. Diptera Brachycera and phona). Type-locality: New York. - Distr.: Athericera of the Fiji Islands ... : 161 NEA: Maryland, New York, Ohio. (Milichia). Type-locality: Lautoka (Fiji).­ flava Sabrosky, 1953. Ent. News 64:38 (Eusi­ Distr.: AUS: Fiji. phona). Type-locality: Kanab (Utah). - apicalis Sabrosky, 1965b. Stuttg. Beitr. Naturk. Distr.: NEA: Utah. 138:1 (Milichia). Type-locality: Marangu mira Coquillett, 1897. U.S. Dep. Agric., Div. Ent., (Tanzania). - Distr.: AFR: Tanzania. Tech. Ser. 7:49. (Eusiphona). Type-loca­ argyrata Hendel, 1'913. Supplta ent. 2:107 lities: Indiana & Colorado. - Distr.: NEA: (Milichia). Type-locality: Kankau, Koshun Washington and Alberta to Prince Edward (Taiwan). - Distr.: OR!: Taiwan. Is., s. to California, Mexico (V.C.), and argyratoides Collin, 1921. Trans. R. ent. Soc. Florida. Lond. 1921:510 (Milichia). Type-locality: vittata Sabrosky, 1982. Revta Mus. La Plata 13:8 Ibadan (Nigeria). - Distr.: AFR: Nigeria. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 103 brevifacialis Deeming, 1976. Ent. scand. 7:38 Iud ens (Wahlberg, 1847). Ofvers. K. VetenskAkad. (Milichia). Type-locality: Ikorodu, Lagos Forh. 4(9): 259 (Lobioptera). Type-locali­ State (Nigeria). - Distr.: AFR: Nigeria. ty: Gusum (Sweden). - Distr. : PAL: widesp. brevirostris (de Meijere, 1909). Tijdschr. Ent. 52: in Europe; former USSR: North European 172 (Prosaetomilichia). Type-locality: tenitory. Djakarta (Java). - Distr.: ORr: Indonesia palposa Zetterstedt, 1848. Dipt. Scand. 7: (Java) 2722 (Milichia). Type-localities: Stock­ cornesi Deeming, 1976. Ent. scand. 7:37 (Milichia). holm, "Abusa", & Uirketorp (Sweden). Type-locality: Agege, Lagos State minuta Walker, 1865. J. Proc. Linn. Soc. Lond. (Nigeria). - Distr.: AFR: Nigeria. Zool. 8: 129 (Milichia). Type-locality: Irian decora (Loew, 1870). Z. ges. Naturw. (N. F. 1) 35: J a ya (Indonesia). - Distr.: A US: Indonesia 9 (Lobioptera). Type-locality: Corsica (Irian Jaya). (France). - Distr.: PAL: France. (Corsican mixta Becker, 1907b. AnnIs hist.-nat. Mus. natn. subspecies ofspeciosa, in Hennig 1937.) hung. 5: 518 (1908a: 160) (Milichia). Type­ dectes Collin, 1921. Trans. R. ent. Soc. Lond. locality: Tenerife (Canary Is.). - Distr.: 1921: 514 (Milichia). Type-locality: Ibadan PAL: Spain (Canary Is.); AFR: South (Nigeria). - Distr.: AFR: Nigeria. Africa. distinctipennis Deeming, 1981. Zool. J. Linn. myrmecophila (de Meijere, 1909). Tijdschr. Ent. Soc. 71: 164 (Milichia). Type-locality: 52: 171 (Prosaetomilichia). Type-locality: Entebbe (Uganda). - Distr.: AFR: Djakarta (Java). - Distr.: ORI: Indonesia Cameroun, Ghana, Ivory Coast, Nigeria, (Java). Rep. Pop. Congo, Uganda. nitida Becker, 1907b. AnnIs hist.-nat. Mus. natn. farquharsoni Collin, 1921. Trans. R. ent. Soc. hung. 5:520 (1908a: 161) (Milichia). Type­ Lond. 1921: 514 (Milichia). Type-locality: localities: Gran Canaria (CanalY Is.). - Ibadan (Nigeria). - Distr.: AFR: Ivory Distr.: Spain (Canary Is.). Coast, Nigeria. orientalis Malloch, 1913b. Insecutorinscit. menstr. formicophila Deeming, 1976. Ent. scand. 7: 39 1: 109 (Milichia). Type-locality: Guam (Milichia). Type-locality: Ilaro Forest, W. (Mariana Is.). - Distr.: AUS: Belau, State (Nigeria). - Distr.: AFR: Nigeria. Hawaiian Is., Gilbert Is., Mariana Is., Tonga, fumicostata Deeming, 1981. Zool. J. Linn. Soc. Vanuatu. 71: 167 (Milichia). Type-locality: Cholo patrizii Hennig, 1952. Beitr. Ent. 2:612 (Milichia). (Malawi). - Distr.: AFR: Mali, Tanzania. Type-locality: Nairobi (Kenya). - Distr.: gigantea Deeming, 1979. Bull. Soc. ent. Fr. 84(3- AFR: Kenya, South Africa. 4): 97 (Milichia). Type-locality: LaMaboke piscivora Malloch, 1926. Proc. Linn. Soc. N.S.W. (Central African Rep.). - Distr.: AFR: 51:547 (Milichia). Type-locality: Towns­ Central African Rep. ville (Queensland). - Distr.: AUS: New ilaroensis Deeming, 1976. Ent. scand. 7: 38 South Wales, Queensland. (Milichia). Type-locality: Ilaro Forest, W. posticata Becker, 1907b. AnnIs hist.-nat. Mus. State (Nigeria). - Distr.: AFR: Nigeria. natn. hung. 5:520 (Milichia). Type-locali­ integra Becker, 1922. Denkschr. Akad. Wiss., ties: Samarang & Pasuman (Java). - Distr.: Wi en (Kl. math. - naturw.) 98(1923): 71 ORI: Indonesia (Java). (Milichia). Type-localities: Tonga & EI proectes Collin, 1921. Trans. R. ent. Soc. Lond. Obeid (Sudan). - Distr.: AFR: ?Egypt, 1921 :512 (Milichia). Type-locality: Ibadan ?Rhodesia, Sudan. (Nigeria). - Distr.: AFR: Nigeria. 104 Irina Brake ENT. SCAND. SUPPL. 57 (2000) prosaetes Collin, 1921. Trans. R. ent. Soc. Lond. Seychelles. 1921 :513 (Milichia). Type-locality: Ibadan sylvicola Deeming, 1981. Zoo!. J. Linn. Soc. 71: 168 (Nigeria). - Distr.: AFR: Nigeria. (Milichia). Type-locality: Agege, Lagos pseudoludens Papp, 1977. Actazool. hung. 23: 173 State (Nigeria). - Distr.: AFR: Nigeria. (Milichia). Type-locality: Szada(Hungary). - Distr.: PAL: Hungary. Milichiella Giglio-Tos, 1895 pubescens Becker, 1907b. AnnIs hist.-nat. Mus. natn. hung. 5:519 (1908a:162) (Milichia). Giglio-Tos, 1895. AnnIs Soc. ent. Fr. 64:367. Type-locality: Gran Canaria (Canary Is.).­ Type-species: Tephritis argentea Fabricius, Distr.: PAL: Egypt, Greece, Israel, Spain 1805. Syst. antl.: 323 (mon.; misidentifica­ (Canary Is.); AFR: Cape V erdeIs., Nigeria, tion ofthe species which was subsequently Oman, Somalia, South Africa, Sudan, described as Milichiella tosi Becker, 1907; Yemen; OR!: Pakistan, ?Taiwan. see Sabrosky 1973). mediocris Sabrosky, 1958b. Stuttg. Beitr. Ophthalmomyia Williston, 1896. Trans. R. ent. Naturk. 4: 1 (Milichia). Type-locality: Soc. Lond. 1896:426. Type-species: Lobio­ Mogadishu (Somalia). ptera lacteipennis Loew, 1866. Bed. ent. savannaticola Deeming, 1981. Zool. J. Linn. Soc. Z. 9(1865): 185 (mon.). 71:167 (Milichia). Type-locality: Samaru, Stenoporomyia Hendel, 1903. Wien. ent. Ztg 22: Zaria (Nigeria). - Distr.: AFR: Nigeria. 250. Type-species: Lobioptera tiefii Mik, seguyi Hennig, 1937. 60a. Milichiidae et Camidae, 1887. Verh. zool.-bot. Ges. Wi en 37:178 Fliegen pal. Reg. 6(1):23 (Milichia). Type­ (orig. des.). (= argyrogaster (Perris, 1876)). locality: Yakouren (Algeria). - Distr. : PAL: Eccoptomma Becker, 1907b. AnnIs hist.-nat. Mus. Algeria. natn. hung. 5 :540. Type-species: Eccoptom­ sinaiensis Steyskal, 1966. Bull. Soc. ent. Egypte majl-ontale Becker, 1907b. l.c.: 542 (des. 109: 120 (Milichia). Type-locality: South Malloch 1934. Dipt. Patag. & S. Chile4(5): Sinai (Egypt). - Distr.: AFR: Egypt, Israel, 463). ?South Africa. Pareccoptomma Duda, 1935b. Stylops 4:25 (as speciosa Meigen, 1830. Syst. Beschr. 6: 132 (Mili­ subgenus ofMilichia). Type-species: Mili­ chia). Type-localities: Marseille & chia nigeriae Duda, 1935b. l.c.: 26 (orig. "Provence" (France). - Distr.: PAL: widesp. des.). in Europe s. of Czech Rep. , Israel, Morocco; former USSR: ?Far East, South European aberrata Becker, 1907b. AnnIs hist.-nat. Mus. territory. natn. hung. 5:538 (Milichiella). Type­ marginata (Mik, 1864). Verh. zool.-bot. Ges. localities: Sorata (Bolivia) & Rosalina Wien 14: 796 (Lobioptera). - Type-locality: (Peru). - Distr.: NEO: Bolivia, Peru. "Rubbia". acrosticalis (Sabrosky, 1958b). Stuttg. Beitr. speciosa canariensis Becker, 1907b. AnnIs hist.­ Naturk. 4:2 (Eccoptomma). Type-locality: nat. Mus. natn. hung. 5:517 (Milichia). Ngaruka, nr. Meru (Tanzania). - Distr.: Type-localities: La Palma & Gran Canaria AFR: Kenya, Nigeria, Tanzania. (Canary Is.). - Distr.: PAL: Spain (Canary arcuata (Loew, 1876). Z. ges. Naturw. 1876:339 Is.). (Lobioptera). Type-locality: Long Island superba Lamb, 1914. Trans. Linn. Soc. Lond. (2, (New York). - Distr.: NEA: New York. Zool.) 16:359 (Milichia). Type-locality: argentea (Fabricius, 1805). Syst. antI.: 323 Mahe (Seychelles). - Distr.: AFR: (Tephritis). Type-locality: Amboina, ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 105

Moluccas (Indonesia). - Distr.: AUS: In­ Distr.: NEO: Peru. donesia (Moluccas), ?Solomon Is. circularis Aldrich, 1931. Proc. Hawaii. ent. Soc. argenteocincta Johnson, 1919. Bull. Am. Mus. 7: 397 (Milichiella). Type-locality: Hawaii. nat. Hist. 41:449 (Milichiella). Type-loca­ - Distr.: AUS: Hawaii. lity: Kingston (Jamaica). - Distr.: NEO: circularis Illingworth, 1929. Proc. Hawaii. West Indies (Jamaica). ent. Soc. 7:234 (Nomen nudum). argentiventris Hendel, 1931. Bull. Soc. ent. Egypte concavum (Becker, 1907b). Annis hist.-nat. Mus. (N.S.) 15(2):72 (Milichiella). Type-locality: natn. hung. 5:541 (Eccoptomma). Type­ Wadi Kanssisrob, Gebel Elba (Egypt). - locality: Santiago (Chile). - Distr.: NEO: Distr.: AFR: ?Cape Verde Is., Egypt, ?Israel, Chile. Nigeria, Oman, Sudan, Yemen. dimidiata (Wiedemann, 1830). Aussereuropiiische argyrogaster (Perris, 1876). Annis Soc. entFr. (5) zweifhigelige Insekten 2:597 (). 6:208 (Milichia). Type-locality: "Departe­ Type-locality: Cordua (Surinam). - Distr.: ment Landes" (France). - Distr.: PAL: NEO: Surinam. Austria, Czech Rep., France, Hungary, flavipalpis Hendel, 1932. Konowia 11: 142 (Mili­ Slovakia; former USSR: North European chiella). Type-locality: San Jose de territory, Far East (Ussuri region). Chiquitos (Bolivia). - Distr.: NEO: Argen­ tiejii (Mik, 1887). Verh. zool.-bot. Ges. Wi en tina, Bolivia. 37: 178 (Lobioptera). Type-locality: frontale (Becker, 1907b). Annis hist.-nat. Mus. Villach, Kiirnten (Austria). natn. hung. 5:542 (Eccoptomma). Type­ bakeri Aldrich, 1931. Proc. Hawaii. ent. Soc. 7: locality: Coronel (Chile). - Distr.: NEO: 398 (Milichiella). Type-locality: Mt. Maki­ Chile. ling, Luzon (Philippines). - Distr.: ORI: hen deli Brake, n.n. for nitida Hendel, 1911. - Philippines, Taiwan. Distr.: NEA: California. bimaculata Becker, 1907b. Annis hist.-nat. Mus. nitida Hendel, 1911. Wien. ent. Ztg 30:39 natn. hung. 5:534 (1908a: 163) (Milichiella). (Milichiella). Type-locality: Pacific Grove Type-locality: Gran Canaria (Canary Is.).­ (California). [Preocc. Walker, 1836]. Distr.: PAL: Spain (Canary Is.). javana de Meijere, 1911. Tijdschr. Ent. 54:427 jreyi (Hendel) in Frey 1936. Commentat. bioI. (Milichiella). Type-locality: Semarang 6: 125 (Eccoptomma). Type-locality: Tene­ (Java). - Distr.: OR!: Indonesia (Java). rife (Canary Is.). lacteipennis (Loew, 1866). Berl. ent. Z. 9(1865): bisignata Melander, 1913a. JI. N.Y. ent. Soc. 21: 185 (Lobioptera). Type-locality: Cuba. - 239 (Milichiella). Type-locality: Riverton Distr. : PAL: Afghanistan, Israel, Marocco, (New Jersey). - Distr.: NEA: Michigan to Spain (Canary and Madeira Is.); almost New Jersey, s. to Kansas and North Caro­ cosmopolitan. lina, also Mexico, New Mexico; NEO: Costa albohalterata (Seguy, 1933). Mems Estud. Rica. Mus. zool. Univ. Coimbra67:50 (Milichia). cinerea (Coquillett, 1900). Proc. U.S. natn. Mus. (Nomen nudum, attributed to Becker acc. 22:268 (Ophthalmomyia). Type-locality: Sabrosky 1980). Bayamon (Puerto Rico). - Distr.: NEO: nigrella Cole, 1912. First Annu. Rep. Laguna. i

West Indies (Puerto Rico). Mar. Lab. 1912:162. Type-locality: Cali- I. cingulata Becker, 1907b. Annis hist.-nat. Mus. fornia. (not synonymous acc. Cole 1969). natn. hung. 5:538 (Milichiella). Type-loca­ lacteiventris Malloch, 1931. Proc. Linn. Soc. N. lity: Pichis, Puelio Bermudes (Peru). - S.W. 56:77 (Milichiella). Type-locality: 106 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Eidsvold (Queensland). - Distr.: AUS: Type-locality: Chiramba (Mozambique). - ?Mariana Is, Queensland. Distr.: Africa: Mozambique. longiseta Hardy & Delfinado, 1980. Insects of smithi Aldrich, 1931. Proc. Hawaii. ent. Soc. 7 :399 Hawaii 13:365 (Milichiella). Type-locality: (Milichiella). Type-locality: Bangkok Honolulu, Oahu (Hawaii). - Distr.: AUS: (Thailand). - Distr.: ORI: Thailand. Hawaiian Is. solitaria (Lamb, 1914). Trans. Linn. Soc. Lond. lucidula Becker, 1907b. AnnIs hist.-nat. Mus. 16(2):361 (Eccoptomma). Type-locality: natn. hung. 5:537 (Milichiella). Type­ Mahe (Seychelles). - Distr.: AFR: Sey­ locality: Callanga (Peru). - Distr.: NEO: chelles, ?U ganda. ? Argentina, Peru; NEA: ?Illinois. spinthera Hendel, 1913. Supplta ent. 2: 107 melaleuca (Loew, 1863). Wien. ent. Mschr. 7:16 (Milichiella). Type-locality: Pilam (Argyrites). Type-locality: "Bloemfontein" (Taiwan). - Distr.: ORI: Taiwan. (South Africa). - Distr.: AFR: South Africa, sumptuosa de Meijere, 1911. Tijdschr. Ent. 54:426 ?Zambia. (Milichiella). Type-locality: Semarang argentea (misidentifications) authors, not (Java). - Distr.: ORI: Indonesia (Java), Fabricius (acc. Sabrosky 1980). Nepal. tosi Becker, 1907b. AnnIs hist. hat. Mus. natn. tricincta Becker, 1907b. AnnIs hist.-nat. Mus. hung. 5:536 (Milichiella). Type-locality: natn. hung. 5: 540 (Milichiella). Type­ Seychelles. locality: Arica (Chile). - Distr.: NEO: Chile. montanum (Becker, 1907b). AnnIs hist.-nat. Mus. tristis Lamb, 1914. Trans. Linn. Soc. Lond. 16(2): natn. hung. 5: 541 (Eccoptomma). Type­ 360 (Milichiella). Type-locality: Mahe locality: Cuzco (Peru). - Distr.: NF(): Chile, (Seychelles). - Distr.: AFR: Seychelles, Bolivia, Peru. Kenya. nigeriae (Duda, 1935b). Stylops 4: 26 (Milichia). unicolor (de Meijere, 1906). Z. syst. Hymenopt. Type-locality: Gadau (Nigeria). - Distr.: Dipterol. 6:333 (Milichia). Type-locality: AFR: Nigelia. Victoria (Cameroun). - Distr.: AFR: nigripes Malloch, 1931. Proc. Linn. Soc. N.S.W. Cameroun; ORI: ?Indonesia (Java). 56: 77 (Milichiella). Type-locality: "not urbana Malloch, 1913c. Can. Ent. 45:284 (Mili­ given" (Australia). - Distr.: AUS: New chiella). Type-locality: Washington (Dis­ South Wales. trict ofColumbia ). - Distr.: NEA: Delaware, nitida(Walker, 1836). Trans. Linn. Soc. Lond. 17: Distr. of Columbia, Michigan, Pennsylva­ 359 (Gymnopa). Type-locality: St. Cathe­ nia, Tennessee, Texas. rine (Brazil). - Distr.: NEO: Brazil. [Comb. populi Steyska1, 1949. Pap. Mich. Acad. Sci., Sabrosky 1973, giving rise to the secondary Ann Arbor 35:130 (Milichiella). Type­ homonymy of nitida Hendel, 1911]. locality: Grosse Ile (Michigan). nudiventris Becker, 1907b. AnnIs hist.-nat. Mus. velutina Becker, 1907b. AnnIs hist.-nat. Mus. natn. hung. 5:537 (Milichiella). Type­ natn. hung. 5:539 (Milichiella). Type­ locality: Mapiri, Sarampiuni (Bolivia). - locality: Rosalina (Peru). - Distr.:NEO: Distr.: NEO: Bolivia. Peru. parva (Macquart, 1843). Mem. Soc. Sci. Agric. vidua Becker, 1907b. AnnIs hist.-nat. Mus. natn. Lille 1842:426 (Milichia). Type-locality: hung. 5:539 (Milichiella). Type-locality: Reunion. - Distr.: AFR: France (Reunion). Umahuankiali (Peru). - Distr.: NEO: Peru. pseudodectes (Seguy, 1933). Mems Estud. Mus. zool. Univ. Coimbra 67:50 (Milichia). ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 107

Pholeomyia Bilimek, 1867 argyrogastra (error) Hendel1932. Konowia 11:135. Bilimek, 1867. Verh. zool.-bot. Ges. Wien 17:903. comans Sabrosky, 1959. Ann. ent. Soc. Am. 52:330 Type-species: Pholeomyia leucozona Bilimek, (Pholeomyia). Type-locality: Flatwoods, 1867. l.c.: 903 (mon.) Rapides Parish (Louisiana). - Distr.: NEA: Rhynchomilichia Hendel, 1903. Wien. ent. Ztg 22: Louisiana. 250. Type-species: Lobioptera argyro­ dampfiSabrosky, 1959. Ann. ent. Soc. Am. 52:328 phenga Schiner, 1868. Reise del' 6ster­ (Ph 0 leomyia ). Type-locality: Peten (Guate­ reichischen Fregatte Novara 2 (Abt.1, Sec. mala). - Distr.: NEO: Guatemala, Mexico. B): 291 (orig. des.; misidentification, de­ decorior Steyskal, 1943. Ent. News 54: 100 scribed as R. schineri n. sp. by Hendel, (Pholeomyia). Type-locality: Orton PI., 1932). Brunswick County (North Carolina). - Pseudomilichia Becker, 1907b. AnnIs hist.-nat. Distr.: NEA: New Jersey to Florida. Mus. natn. hung. 5:542. Type-species: dispar (Becker, 1907b). AnnIs hist.-nat. Mus. natn. Pseudomilichia schnusei Becker, 1907b. hung. 5:524 (Rhynchomilichia; as leuco­ l.c.: 543 (des. by Hendel 1932. Konowia gastra val'.). Type-locality: Tifton 11: 135). (Georgia). - Distr.: NEA: Alabama, Florida, Paramilichia Malloch, 1913a. Proc. U.S. natn. Georgia, Texas, Virginia. Mus. 46: 135. Type-species: Milichia excelsior (Becker, 1907b). AnnIs hist.-nat. Mus. longiseta Becker, 1907b. AnnIs hist.-nat. natn. hung. 5: 526 (Rhynchomilichia). Type­ Mus. natn. hung. 5:530 (mon.). locality: Mapiri, Sarampiuni (Bolivia). - Macromilichia Hendel, 1932. Konowia 11:140. Distr.: NEO: Bolivia. Type-species: Macromilichia nigricosta expansaAldrich, 1925. Proc. U.S. natn. Mus. 66: 1 Hendel, 1932. Konowia 11:l42(orig. des.). (Pholeomyia). Type-locality: Mount Lowe (California). - Distr.: NEA: California. aequatorialis Seguy, 1934. RevtaSoc. ent. argent. fasciventris (Becker, 1907b). AnnIs hist.-nat. Mus. 6: 14 (Pholeomyia). Type-locality: EINapo natn. hung. 5:528 (Rhynchomilichia). Type­ (Ecuador). - Distl'.: NEO: Ecuador. locality: Mapiri, Sarampiuni (Bolivia). - anomalaHendel, 1933b. RevtaEnt.,Rio del 3:82 Distr.: NEO: Bolivia. (Pholeomyia). Type-locality: Soledade hurdi Sabrosky, 1959. Ann. ent. Soc. Am. 52:324 (Brazil). - Distr.: NEO: Brazil. (Pholeomyia). Type-locality: Ahuacatlan, anthracina (Becker, 1907b). AnnIs hist.-nat. Mus. Nayarit (Mexico). - Distr.: NEO: Mexico. natn. hung. 5:525 (Rhynchomilichia). Type­ implicata (Becker, 1907b). AnnIs hist.-nat. Mus. locality: Asuncion (Paraguay). - Distr.: natn. hung. 5: 543 (Pseudomilichia). Type­ NEO: Paraguay. locality: Pichis, Puelio Berrnudes (Peru).­ argyrata Hendel, 1932. Konowia 11:137 (Phole­ Distr.: NEO: Peru. omyia). Type-locality: Tapikiole (Argenti­ indecora (Loew, 1869). Berl. ent. Z. 13: 50 na). - Distr.: NEO: Argentina. (Lobioptera). Type-locality: Nebraska. - argyrophenga (Schiner, 1868). Reise der oster­ Distr.: NEA: British Columbia to Nova reichischen Fregatte Novara 2 (Abt.1, Sec. Scotia, s. to California, Kansas, and Georgia. B): 291 (Lobioptera). Type-locality: "South insecta (Becker, 1907b). AnnIs hist.-nat. Mus. America". - Distr.: NEO: Bolivia, Peru. natn. hung. 5:527 (Rhynchomilichia). Type­ argyrocephala (error) Hendel 1932. Konowia locality; Mapiri, S. Carlos (Bolivia). - 11:135. Distr.: NEO: Bolivia. 108 Irina Brake ENT. SCAND. SUPPL. 57 (2000) latifrons Sabrosky, 1959. Ann. ent. Soc. Am. 52: pectoralis Hendel, 1932. Konowia 11: 138 (Pholeo­ 321 (Pholeomyia). Type-locality: Gaver­ myia). Type-locality: Villa Montes (Boli­ nor's Harbor, Eleuthera Is. (Bahama Is.).­ via). - Distr.: NEO: Bolivia. Distr.: NEO: West Indies (Bahama Is., politifacies Sabrosky, 1959. Ann. ent. Soc. Am. ?Cuba). 52:323 (Pholeomyia). Type-locality: Cam­ leucogastra (Loew, 1861). Wien. ent. Mschr. 5(2): bito, Provo Trujillo (Dominican Rep.). - 43 (Milichia). Type-locality: Cuba. - Distr.: Distr.: NEO: Brazil, Colombia, Costa Rica, NEO: Mexico, West Indies (Cuba, st. El Salvador, Mexico, Panama Canal Zone, Vincent). West Indies (Barbados, Dominican Rep., leucozona Bilimek, 1867. Verh. zool.-bot. Ges. Puerto Rico, Trinidad); AUS: Society Is. Wi en 17:903 (Pholeomyia). Type-locality: (Tahiti); AFR: Cameroon, Ghana, Ivory Cave Cacahuamilpa (Mexico). - Distr.: Coast, Nigeria. NEO: Guatemala, Mexico. praeocellaris Hendel, 1932. Konowia 11: 139 cacahuamilpensis Herrera, 1892. Estudio 4: (Pholeomyia). Type-locality: Villa Montes 272 (Pholeomyia) (unnecessary n.n.). (Bolivia). - Distr.: NEO: Bolivia. longifacies Hendel, 1933b. RevtaEnt., Rio de J. 3: praesecta (Becker, 1907b). AnnIs hist.-nat. Mus. 81 (Pholeomyia). Type-locality: "Brazil" natn. hung. 5:525 (Rhynchomilichia). Type­ & San Bernardino (Paraguay). - Distr.: locality: Mouth of Rio Pachitea (Peru). - NEO: Brazil, Paraguay. Distr.: NEO: Peru. longiseta (Becker, 1907b). AnnIs hist.-nat. Mus. prominens (Becker, 1907b). AnnIs hist.-nat. Mus. natn. hung. 5:530 (Milichia). Type-locality: natn. hung. 5 :526 (Rhynchomilichia). Type­ Asuncion (Paraguay). - Distr.: NEO: localities: Chanchamayo (Peru) & Mapiri, Argentina, Nicaragua, Paraguay. Serampiuni (Bolivia). - Distr.: NEO: myopaMelander, 1913a. JI. N.Y. ent. Soc. 21:238 Bolivia, Peru. (Pholeomyia). Type-locality: Haiti. - Distr.: pseudodecora (Becker, 1907b). AnnIs hist.-nat. NEO: Bolivia, Brazil, Costa Rica, Mexico, Mus. natn. hung. 5:524 (Rhynchomilichia). Panama Canal Zone, Venezuela, West Type-locality: Tifton (Georgia). - Distr.: Indies (Bahama Is., Cuba, Dominica, NEA: Florida, Georgia, Maryland, South Dominican Rep., Haiti, Puerto Rico, Carolina. Trinidad); NEA: Florida. quadrifasciata Hendel, 1932. Konowia 11:139 nigricosta (Hendel, 1932). Konowia 11: 142 (Pholeomyia). Type-locality: Esmeralda, (Macromilichia). Type-locality: "EI Cairo", Rio Pilcomayo (Bolivia). - Distr.: NEO: Santa Cruz de la Sierra (Bolivia). - Distr.: Bolivia. NEO: Bolivia. robertsoni (Coquillett, 1902). JI. N.Y. ent. Soc. nitidula Sabrosky, 1959. Ann. ent. Soc. Am. 52:327 10: 187 (Milichia). Type-locality: Inverness (Pholeomyia). Type-locality: Georgia. - (Florida). - Distr.: NEA: Florida. Distr.: NEA: Georgia. schineri Hendel, 1932. Konowia 11: 136 obscuraSabrosky, 1959.Ann. ent. Soc. Am. 52:330 (Pholeomyia). Type-locality: Brazil. - (Pholeomyia). Type-locality: Austin Distr.: NEO: Argentina, Brazil, Paraguay. (Texas). - Distr.: NEA: Texas. argyrophenga (misidentifications) Hendel palparis (Becker, 1907b). AnnIs hist.-nat. Mus. 1903 and Becker 1907 (acc. Hendel 1932). natn. hung. 5: 524 (Rhynchomilichia). Type­ schnusei (Becker, 1907b). AnnIs hist.-nat. Mus. locality: Asuncion (Paraguay). - Distr.: natn. hung. 5:543 (Pseudomilichia). Type­ NEO: Paraguay. localities: Meshagua (Peru) & Mapiri, San ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 109

Antonio (Bolivia). - Distr.: NEO: Bolivia, zoologischen Expedition nach dem anglo-iigyptischen Peru. Sudan (Kordofan) 1914. VI. Diptera. - Denkschr. Akad. Wiss., Wien 98 (1923):57-82. sororcula (Becker, 1907b). AnnIs hist.-nat. Mus. Bezzi, M. 1928. Diptera Braehycera and Athericera of the natn. hung. 5 :521 (Rhynchomilichia). Type­ Fiji Islands based on material in the British Museum (Natural History). British Museum (Nat. Hist.), London: localities: Bolivia & Peru. - Distr.: NEO: viii + 220 pp. Bolivia, Peru. Bilimek, D. 1867. FaunaderGrotte CacahuamilpainMexiko. texensis Sabrosky, 1959. Ann. ent. Soc. Am. 52:329 - Verh. zool.-bot. Ges. Wi en 17:901-908. Brunetti, E. 1924. Tipulidae, Tabanidae, , (Pholeomyia). Type-locality: San Antonio Acalyptratae, Muscidae and Phoridae. 111: Brunetti, E., (Texas). - Distr.: NEA: Texas. Edwards, F. W., Lamb, C. G. & Patton, W. S. (Eds): vockerothi Sabrosky, 1961. Ent. News 72:231 Diplera of the Siju Cave, Garo Hills, Assam. - Rec. Indian Mus. 26:99-112, 5 figs. (Pholeomyia). Type-locality: Highlands Chandler, P. J. 1998. Milichiidae. Pp. 143-144 in: Chandler, (NOlih Carolina). - Distr.: NEA: North P. J. (Ed): Checklists of insects of the British Isles (New Carolina. Series), Part 1: Diptera (IncOlporating a list of Irish Diptera). Handbooks for the indentification of British Ulia Becker, 1907 insects 12:xx + 234 pp. Cole, F. R. 1912. Some Diptera of Laguna Beach. - First Becker, 1907b. AnnIs hist.-nat. Mus. natn. hung. 5: Annu. Rep. Laguna Mar. Lab. 1912:150-162. 544. Cole, F. R. 1969. Milichiidae. In: The flies of Western North Type-species: Ulia poecilogastra Becker, America. University of California Press, Berkeley, pp. 387-390. 1907b. l.c.: 544 (mon.). Collin, J. E. 1921. Description of a new genus and two new species of Cecidomyidae, and six new species of poecilogastra Becker, 1907b. AnnIs hist. -nat. Mus. Acalyptrate Muscidae (Ephydridae and Milichiidae).­ natn. hung. 5:544 (Ulia). Type-localities: Trans. R. ent. Soc. Lond. 1921:504-517. Coquillett, D. W. 1897. Revision of the of Pichis & Rosalina (Peru). - Distr.: NEO: America north of Mexico. - U.S. Dep. Agric., Div. Bolivia, Costa Rica, Peru. Ent., Tech. Ser 7: 154 pp. Coquillett, D. W. 1900. Report on a collection of dipterous insects from Puerto Rico. - Proc. U.S. natn. Mus. References 22:249-270. Coquillett, D. W. 1902. New North American Diptera. - JI. Aldrich, J. M. 1925. New Diptera or two-winged flies in the N.Y. ent. Soc. 10:186-191. United States National Museum. - Proc. U.S. natn. Curran, C. H. 1927. A new species of Mallochiella (Diptera: Mus. 66:1-36. Milichiidae). - Can. Ent. 59:49-50. Aldrich, J. M. 1926. Descriptions of new and little-known Cunan, C. H. 1930. Report on the Diptera collected at the Diptera or two-winged flies. - Proc. U.S. natn. Mus. station for the study of insects, Haniman Interstate Park, 69:1-26. N.Y. -Bull. Am. Mus. nat. Hist. 61 (1931):21-115. Aldrich, J. M. 1931. New acalyptrate Diptera from the Curran, C. H. 1934. The families and genera of North Pacific and Oriental Regions. - Proc. Hawaii. ent. Soc. American Diptera. 512 pp, 235 figs, 2 pIs. New York. 7:395-399. Curran, C. H. 1936. The Templeton Crocker Expedition to Becker, T. 1903. Agyptische Dipteren (Fortsetzung und the western Polynesian and Melanesian Islands, 1933. Schlu/3). - Mitt. zool. Mus. Berl. 2:67-195. - Proc. Calif. Acad. Sci. (4) 22:1-66. Becker, T. 1907a. Desmometopa. - Wien. ent. Ztg 26: 1-5. Deeming, J. C. 1976. Four new short-faced species of the Becker, T. 1907b. Die Dipteren-Gruppe Milichiinae. - genus Milichia Meigen with a key to the Nigerian AnnIs hist.-nat. Mus. natn. hung. 5:507-550. members of the group (Diptera: Milichiidae). - Ent. Becker, T. 1908a. Dipteren der Kanarischen Inseln. - Mitt. scand. 7:35-40. zool. Mus. Berl. 4:1-180. Deeming, J. C. 1979. A new giant species of Milichia Becker, T. 1908b. Diptera duo nova ex Hungaria. - AnnIs Meigen from Central Africa. - Bull. Soc. ent. Fr. hist.-nat. Mus. natn. hung. 6:320. 84:97-99. Becker, T. 1910. Dipterologische Sammelreise nach Korsika. Deeming, J. C. 1981. Some new AfricanMilichiidae (Diptera: - Dt. ent. Z. 1910:635-665. Cyc1orrhapha) having pronounced sexual dimorphism Becker, T. 1922. Wissenschaftliche Ergebnisse der mit of the frons. - Zool. J. Linn. Soc. 71:159-169. Unterstiitzung der Akademie der Wissenschaften in Wien Duda, O. 1932.61. Chloropidae. In: Lindner, E. (Ed): Die aus der Erbschaft Treitl von F. Werner unternommenen Fliegen der palaearktischen Region. Vol. 6(1). 248 pp. 110 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

Stuttgart, Schweizerbart. Konowia 11:98-110; 115-145, 15 figs. Duda, O. 1935a. Beitrag zur Kenntnis del' palaarktischen Hendel, F. 1933a. Neue acalyptrate Musciden aus del' pala- Madizinae. - Natuurh. Maandbl. 24: 14-16, 24-26, 37- arktischen Region (Dipt.). - Dt. ent. Z. 1933:39-56. 40. Hendel, F. 1933b. Ober einige Typen Wiedemann's und Duda, O. 1935b. Einige neue afrikanische akalyptrate Schiner's von acalYjJtaten Musciden aus Slidamerika, Musciden (Diptera) des British Museum. - Stylops \ nebst einigen verwancrten Arten (Dipt.). - Revta En!., 4:25-34. Rio de J. 3:58-83, 1 fig. Enderlein, G. 1921. Dipterologische Studien XVII. Zool. Hennig, W. 1937. 60a. Milichiidae et Carnidae. In: Lindner, Anz.52:219-232. E. (Ed):Die Fliegen del' palaearktischen Region. Vol. Enderlein, G. 1927. Dipterologische Studien XIX. Stettin. 6(1). 91 pp. Stuttgart, Schweizerbart. ent. Ztg 88(1):102-109. Hennig, W. 1952. Bemerkenswerte neue Acalyptraten in der Enderlein, G. 1934. Dipterologica. I. - Sber. Ges. naturf. Sammlung des Deutschen Entomologischen Institutes Freunde Berl. 1933:416-429. (Diptera: Acalyptrata). - Beitr. Ent. 2:604-618. Fabricius, J. C. 1805. System a antliatorum secundum ordines, Herrera, A. L. 1892. Fauna cavernicola de Cacahuamilpa. genera, species adiectis synonymis, locis, observationi­ Pp. 268-281, pIs. 14-15 in: Arriaga, J. J., Urbina, M. & bus, descriptionibus. Carolum Reichard. 1-372 + 30 pp. Rebollar, R. (Eds): Expedicion a la gruta Cacahuamilpa. Brunsvigae. - Estudio, Semenario de Cien. Med. (Inst. Med. Nac. de Fallen, C. F. 1810. Specimen entomologicum novam Diptera Mex.) (1893) 4:268-281, 339-344, 455-461, pIs. 14- disponendi methodum exhibens. Litteris Berlingianis. 15,17. 26 pp, 1 Tafel. Lundae. Illingworth, J. F. 1926. Desmometopa m-nigrul1l (Zetterstedt) Fallen, C. F. 1820. Oscinides Sveciae. Vol. 2 (20). Litteris [Note]. - Proc. Hawaii. ent. Soc. 6:224. Berlingianis. II + 10 pp. Lundae. Illingworth,1. F. 1929. Desmometopa tarsalis Loew [Note]. Fallen, C. F. 1823. Phytomyzides et Ochtidiae Sveciae. - Proc. Hawaii. ent. Soc. 7:233-234. Litteris Berlingianis.1 0 pp. Lundae. Iwasa, M. 1997. A new species of the genus Aldrichiomyza Frey, R. 1918. Mitteilungen liberslidamerikanische Dipteren. Hendel (Diptera, Milichiidae) from Japan. - J ap. J. Ent. - Ofvers. finska VetenskSoc. Forh. 60 (1917-1918, 65: 826-829. Afd. A, no. 14):35 pp, 1 pI. Johnson, C. W. 1919. A revised list of the Diptera of Frey, R. 1936. Die Dipterenfauna der Kanarischen Inseln Jamaica. - Bull. Am. Mus. nat. Hist. 41:421-449. und ihre Probleme. - Commentat. bioI. 6(1): 1-237. Kertesz, K. 1897. Dipterologisches aus Neu-Guinea. - Giglio-Tos, E. 1895. Mission scientifique de M. Ch. Alluaud Termeszetr. Fliz. 20:611-613. aux lies Sechelles (Mars-Avril-Mai 1892) 5e memoire. Kertesz, K. 1899. Verzeiclmis einiger, von L. Biro in Neu­ Dipteres. - AnnIs Soc. ent. Fr. 64:353-368. Guinea und am Malayischen Archipe1 gesammelten Hardy, D. E. & Delfinado, M. D. 1980. Family Milichiidae. Dipteren. - Termeszetr. Fliz. 22:173-195. Pp. 351-367 in: Insects of Hawaii 13:451 pp. Kramer, H. 1920. Zwei neue deutsche Musciden. - Zool. Hendel, F. 1903. Kl'itische Bemerkungen zur Systematik del' Jb. Syst. 43:331-332, Fig. A. Muscidae acalypteratae. - Wien. ent. Ztg 22:249-252. Lamb, C. G. 1914. The Percy Siaden Trust expedition to the Hendel, F. 1907. Neue und interessante Dipteren aus dem Indian Ocean in 1905, under the leadership of Mr. J. kaiserlichen Museum in Wien. - Wien. ent. Ztg 26:223- Stanley Gardiner, M. A. Vol. 5. No XV. - Trans. Linn. 245,1 pI. Soc. Lond. (2, Zool.) 16:307-372. Hendel, F. 1911. ObervonProf. J. M. Aldrich erhaltene und Latreille, P. A. 1829. Les Crustaces, les Arachnides et les einige andere amerikanische Dipteren. - Wien. ent. Ztg Insectes, (Vol. 2). In: Cuvier, G. C. L. D. (Ed): Le regne 30:19-46. distribue d 'apres son organisation, pour servir de Hendel, F. 1913. H. Sauter's Formosa-Ausbeute. Acalyptrate base it I'histoire naturelle des animaux, et d'introduction Musciden (Dipt.) II. - Supplta ent. 2:77-112. it I'anatomie comparee. Ed. 2, 5:1-556. Paris. Hendel, F. 1914a. Namensanderung (Dipt.). - Ent. Mitt. Lioy, P. 1863-1864. I ditteri distribuiti secondo un nuovo 3:73. metodo di classificazione naturale. Atti 1st. veneto Sci. Hendel, F. 1914b. H. Sauter's Formosa-Ausbeute. Acalyp­ (3) 9:187-236 (1863), 499-518, 569-604, 719-771, trate Musciden (Dipt.) III. - Supplta ent. 3:90-117, 7 879-910,989-1027, 1087-1126, 1311-1352 (1864); figs. 10:59-84. Hendel, F. 1919. Neues liber Milichiiden (Dipt.). - Ent. Loew, H. 1861. Diptera aliquot in insula Cuba collecta.­ Mitt. 8: 196-200. Wien. ent. Mschr. 5(2):33-43. Hendel, F. 1924. Neue europaische Phyllomyza-Arten (Dipt. Loew, H. 1863. Enumeratio ctipterorum, quae C. Tollin ex Milichiidae). - Dt. ent. Z. 1924:405-408. Africa meridionali (Orangestaat, Bloemfontein) misit. Hendel, F. 1931. Neue iigyptische Dipteren aus del' Gruppe - Wien. ent. Mschr. 7:9-16. der acalyptraten Musciden, gesammeJt von Prof. Loew, H. 1866. Diptera Americae septentrionalis indigena. Efflatoun Bey. - Bull. Soc. ent. Egypte 15(2):59-73. - Berl. ent. Z. 9(1865) (Centuria 6, no. 96):127-186. Hendel, F. 1932. Die Ausbeute der deutschen Chaco­ Loew, H. 1869. Diptera Americae septentrionalis indigena. Expedition. 1925126. Diptera XXX-XXXVI. - Centurio octava. - Berl. ent. Z. 13:1-52. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 111

Loew, H. 1870. Lobioptera speciosa Meig. und decora nov. neuerArten. - Verh. zool.-bot. Ges. Wien 14:791-797, sp. - Z. ges. Naturw. (N. F. I) 35:9-14. Tafel 21, Fig. 5. Loew, H. 1876. Beschreibung neueramerikanischer Dipteren. Mik, J. 1887. Ober Dipteren. I. Drei neue iisterreichische - Z. ges. Naturw. (N. F.) 1876:317-340. Dipteren. - Verh. zool.-bot. Ges. Wi en 37:1-8 [173- Macquart, J. 1843. Dipteres exotiques nouveaux ou peu 180], Tafel IV. connus. - Mem. Soc. Sci. Agric. Lille 1842:162-460. Ozerov, A. L. 1992. Xenophyllomyza deserticola new genus Malloch, J. R. 1913a. A synopsis of the genera of Agromy­ new species from Turkmenistan. - Vestnik Zoo 1. zidae, with descriptions of new genera and species. - 1992(2):18-20. Proc. U.S. natn. Mus. 46:127-154, pis. 4-6. Papp, L. 1976. Milichiidae and Carnidae (Diptera) from Malloch, J. R. 1913b. A new species of Agromyzidae Mongolia. - Acta zool. hung. 22:369-387. (Diptera). - Insecutor inscit. menstr. I: 109-110. Papp, L. 1977. New species and records of Hungarian Malloch,J. R. 1913c. New North American Diptera. -Can. Odiniidae, Milichiidae and Carnidae (Diptera). - Acta Ent. 45:282-284. zool. hung. 23: 171-181. Malloch, J. R. 1914. Formosan Agromyzidae. - Annis Papp, L. 1978. Some cavernicolous Diptera of the Geneva hist.-nat. Mus. natn. hung. 12:306-336, pis. 9-10. Museum. - Revue suisse Zool. 85:99-106. Malloch, J. R. 1924. Notes on Australian Diptera. No iii. Papp, L. 1984a. Family Milichiidae. In: S06s, A. & Papp, L. Proc. Linn. Soc. N.S.W. 49:329-338. (Eds): Catalogue of Palaearctic Diptera 10: 110-123. Malloch, J. R. 1925. Notes on Australian Diptera. No. vi.­ Papp, L. 1984b. Cavernicolous Milichiidae: three new species Proc. Linn. Soc. N.S.W. 50:80-97. from Fiji and Sri Lanka. - Revue suisse Zool. 91 :241- Malloch, J. R. 1926. Notes on Australian Diptera. No. ix.­ 247. Proc. Linn. Soc. N.S.W. 51:545-554. Papp, L. 1993. Three new milichiid species from Hungary. Malloch, J. R. 1927. Notes on Australian Diptera. No. x. - - Annis hist.-nat. Mus. natn. hung. 85: 133-139. Proc. Linn. Soc. N.S.W. 52:1-16. Papp, L. 1998.87. Milichiidae. P. 295 in: Merz, B., Bachli, Malloch, J. R. 1931. Notes on Australian Diptera. No. xxvii. G., Haenni, J.-P. & Gonseth, Y. (Eds.): Diptera - - Proc. Linn. Soc. N.S.W. 56:60-78. Checklist. - Fauna Helvetica 1:369 pp. Malloch, J. R. 1934. Acalyptrata (concluded). Pp. 393-489, Perris, E. 1876. Nouvcllc promenade entomologique. '­ pI. 8, figs. 69-84 in: Diptera of Patagonia and South Annis Soc. ent. Fr. (5) 6: 171-244. Chile, 6(5) [British Mus. (Nat. Hist.)]. London. Rolulcek, J. 1997. Milichiidae. P. 80 in: Chvala, M. (Ed): Meigen, J. W. 1830. Systematische Beschreibung der Check List ofDiptera (Insecta) of the Czech and Slovak bekannten europaischen zweifliigeligen Insekten. Vol. Republics. Karolinum. Charles University Press, 130 6. xi + 401 pp. Hamm. pp. Prague. Meigen, J. W. 1838. Systematische Beschreibung der Roser, C. L. F. von 1840. 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New species and notes on North Dipteren. IX. - Tijdschr. Ent. 57: 137-276, pis. 5-7. American acalyptrate Diptera. - En!. News 69: 169- Meijere, J. C. H. de 1916. Studien liber siidostasiatische 176. Dipteren. XI. - Tijdschr. Ent. 59: 184-213. Sabrosky, C. W. 1958b. East African Milichiidae (Diptera). Meijere, J. C. H. de 1940. Eine neue myrmecophile Phyllo- (Ergebnisse der Deutschen Zoologischen Ostafrika­ 111yza. - Ent. Ber., Amst. 10 (232):222-223. Expedition 1951/52, Grnppe Lindner Stuttgart, Nr. 32). Meijere, J. C. H. de 1946. Zevende Supplement op de - Stuttg. Beitr. Naturk. 4:1-5. Nieuwe Naamlijst van Nederlandsche Diptera van 1898 Sabrosky, C. W. 1959. A revision of the genus Pholeomyia (Eerste Supplement op myne Naamlijst van 1939). - in North America (Diptera: Milichiidae). - Ann. en!. Tijdschr. Ent. 87 (1944): 17-18. Soc. Am. 52:316-331. Melander, A. J. 1913a. A synopsis of the dipterous groups Sabrosky, C. W. 1961. Three new nearctic acalyptrate Agromyzidae, Milichiinae, Ochthiphilinae and Geo­ Diptera. - En!. News 72:229-234, I fig. myzinae. - n. N.Y. ent. Soc. 21:219-273, 283-300, Sabrosky, C. W. 1964. Milichiidae and Chloropidae (Diptera) p1.8. from the Batu Caves, Malaya. - Pacif. Insects 6:308- Melander, A. J. 1913b. Some acalyptrate Muscidae. - 311. Psyche, Camb. 20: 166-169, 1 fig. Sabrosky, C. W. 1965a. Family Milichiidae. Pp. 728-733 in: Mik, J. 1864. Dipterologische Beitrage, I. Beschreibung Stone et al. (Eds): A catalog of the Diptera of America 112 Irina Brake ENT. SCAND. SUPPL. 57 (2000)

North of Mexico. Agric. Handbook 276:iv + 1696 pp. Seguy, E. 1934. Etude sur quelques Muscides de I' Amerique Sabrosky, C. W. 1965b. East African Milichiidae and Latine. - Revta Soc. ent. argent. 6:9-16, 3 figs. Chloropidae (Diptera). - Stuttg. Beitr. Naturk. 138:1- Seguy, E. 1938. Diptera L. Nematocera et Brachycera. Pp. 8. 319-380 ill: Jeannel (Ed): Mission Scientifiquede l'Omo; Sabrosky, C. W. 1973.75. Family Milichiidae. 12 pp in: A Vol. 4 (Zoologie). - Mus. natn. Hist. nat. 8. catalogue of the Diptera of the Americas South of the Steyskal, C. 1942. A new species of Phyl/omyza from United States. Museu de Zoologia, Universidade de Sao Virginia (Diptera: Milichiidae). - Ent. News 53 :84-85. Paulo, Sao Paulo, Brazil. Steyskal, G. C. 1943. A new species of Phoieomyia, with a Sabrosky, C. W. 1977. Family Milichiidae. Pp. 270-274 in: key to the NOlihAmerican species (Diptera, Milichiidae). Delfinado, M. D. & Hardy, D. E. (Eds): A Catalog of the -Ent. News 54:99-102. Diptera of the Oriental Region 3:x + 854 pp. Honolulu. Steyskal, G. C. 1949. The dipterous fauna of tree tmnks.­ Sabrosky, C. W. 1979. Faune entomologique de I'archipel Pap. Mich. Acad. Sci. 35:121-134. des Comores. - Mem. Mus. natn. Hist. nat., Paris, Serie Steyskal, G. C. 1966. Notes and descriptions of Egyptian A, Zool. 109:315-330. acalyptrate Diptera (Ephydridae, Milichiidae, Otitidae). Sabrosky, C. W. 1980.75. Family Milichiidae. Pp. 686-689 -Bull. Soc. en!. Egypte 109:109-125. in: Crosskey, R. W. (Ed.): Catalogue of the Diptera of Strobl, P. G. 1900a. Dipterenfauna von Bosnien, Herzegovina the Afrotropical Region. 1437 pp. London. und Dalmatien. Wiss. Mitt. Bosn. Herzeg. 7:552-670. Sabrosky, C. W. 1982. Una nueva especie de Eusiphona de Strobl, P. G. 1900b. Spanische Dipteren. VIII. Theil. - la Republica Argentina. - Revta Mns. La Plata, n. s. Wien. ent. Ztg 19:1-10 Zoo I. 13:7-9. Villeneuve, J. 1920. Vichyia acyglossa, espece et genre Sabrosky, C. W. 1983. A Synopsis of the world species of nouveaux de la famille des Milichiinae (Dipt. Muscidae). Desmometopa Loew (Diptera, Milichiidae). - Contr. - Bull. Soc. ent. Fr. 1920:69-70, Fig. 1. Am. ent. Inst. 19(8):1-69. Wahlberg, P. F. 1847. Nya slagten af Agromyzidae. - Sabrosky, C. W. 1987. A new species of Leptometopa Gfvers. K. Vetensk.Akad:F6i·h. 4:259-263, Tab. 7, fig. (Diptera, Milichiidae) from Madagascar pollinating 1. Ceropegia (Asclepiadaccac). - Proc. ent. Soc. Wash. Walker,F. 1836. Descriptions, etc., of the Diptera. Pp. 331- 89:242-243. 359 in: Curtis, J., Haliday, A. H. & Walker, F. (Eds): Sabrosky, C. W. 1989.78. Family: Milichiidae. In: Evenhuis, Descriptions, etc., of the insects collected by Captain P. N. (Ed): Catalog of the Diptera of the Australasian and P. King, R. N., F. R. S., in the survey of the Straits of Oceanian Regions. Bishop. Mus. Spec. Publ. 86:555- Magellan. - Trans. Linn. Soc. Lond.(2, Zool.) 17:315- 557. 359. Schiner, J. R. 1868. Diptera (Art. 1). In: Reise der Walker, F. 1865. Descriptions ofnew species ofthedipterous ostelTeichischen Fregatte Novara 2 (Abt. 1, Sec. B). 388 insects of New Guinea. -J. Proc. Linn. Soc. 8: 109-130. pp, 4 pIs. Wien. Westwood, J. 0.1840. Diptera. Pp. 125-158 in: Synopsis of Schmitz, S. J. 1923. Drei neue europaische Phyllomyza­ the genera of British insects, in: An introduction to the Arten. - Konowia 2:44-47. modem classification of insects; founded on the natural Schmitz, S. J. 1924. Een nieuwe vlieg uit Zuid-Limburg. - habits and corresponding organization of the different Natuurh. Maandbl. 13:16. families. 2:xi + 1-557. London. Seguy, E. 1932. Spedizione scientifica all'oasi di Cufra Wiedemann, C. R. W. 1830. Anssereuropaische zweifliige­ (Marzo-Luglio 1931). Insectes dipteres. - Annali Mus. lige Insekten. Vol. 2. xii + 684 pp. Aachen. civ. Stor. nat. Giacomo Doria 55(1930-1931):490-511, Williston, S. W. 1896. On the Diptera of St. Vincent (West figs 1-3. Indies). Trans. ent. Soc. Lond. 1896:253-446, pIs. 8-14. Seguy, E 1933. Contributions it I'etude de la faune du Williston, S. W. 1897. Diptera Brasiliana, Part 4. Kans. Mozambique. Voyage de M.M. Lesne (1928-1929) 13e Univ. Q. (A) 6:1-12. note. Dipteres (2e partie). - Mems Estud. Mus. zool. Zetterstedt, J. W. 1848. Diptera scandinaviae disposita et Univ. Coimbra 67:5-80. descripta. 7:2581-2934. Lundae. ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 113

Index abenata Becker, 1907, Milichiella...... 104 acrosticalis (Sabrosky, 1958), (Eccoptomma), Milichiella ...... 104 acyglossa (Villeneuve, 1920), (Vichyia), Neophyllomyza ...... 93 aczeli Sabrosky, 1983, Desmometopa ...... 97 aelleni Papp, 1978, Leptometopa ...... 100 aelleni Papp, 1984, Phyllomyza ...... 94 aequatorialis Seguy, 1934, Pholeomyia ...... 107 aethiops Malloch, 1913, Milichia ...... 102 agromyzina (Hendel, 1911), (Aldrichiella), Aldrichiomyza ...... 93 albipennis (Lamb, 1914), (Madiza), Leptometopa ...... 100 albipennis (Malloch, 1924), (Hypaspistomyia), Leptometopa ...... 100 albohalterata (Seguy, 1933), (Milichia), Milichiella ...... 105 albomaculata (Strobl, 1900), (Lobioptera), Milichia ...... 102 aldabrae Sabrosky, 1983, Desmometopa ...... 97 Aldrichiella Hendel, 1911, Aldrichiomyza ...... 93 Aldrichiomyza Hendel, 1914 ...... 93 angustifrons Bezzi, 1928, Milichia ...... 102 annulimana (von Roser, 1840), (Agromyza), Leptometopa ...... 100 annulitarsis (von Roser, 1840), (Agromyza), Leptometopa ...... 100 annulitarsis (Zetterstedt, 1848), (Madiza), Leptometopa ...... 100 anomala Hendel, 1933, Pholeomyia ...... 107 anthracina (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 107 anuda (Cunan, 1936), (Desmometopa), Neophyllomyza ...... 93 apicalis Sabrosky, 1965, Milichia ...... 102 approximata (Malloch, 1913), (Phyllomyza), Neophyllomyza ...... 94 approximatonervis (Lamb, 1914), (Desmometopa), Neophyllomyza ...... 93 Arctobiella (Coquillett, 1902), Dasiops ...... -> arcuata (Loew, 1876), (Lobioptera), Milichiella ...... 104 argentea (Fabricius, 1805), (Tephritis), Milichiella ...... 105 argentea, authors, not Fabricius, Milichiella ...... 106 argenteocincta Johnson, 1919, Milichiella ...... 105 argentinica Sabrosky, 1983, Desmometopa ...... 97 argentiventris Hendel, 1931, Milichiella ...... 105 argyrata Hendel, 1913, Milichia ...... 102 argyrata Hendel, 1932, Pholeomyia ...... 107 argyratoides Collin, 1921, Milichia ...... 102 Argyritis Latreille in Cuvier, 1829, Milichia ...... 102 argyrocephala (enor) Hendel, 1932, Pholeomyia ...... 107 argyrogaster (PelTis, 1876), (Milichia), Milichiella ...... 105 argyrogastra (enor) Hendel, 1932, Pholeomyia ...... 107 argyrophenga (Schiner, 1868), (Lobioptera), Pholeomyia ...... 107 argyrophenga Schiner of Hendel, 1903, and Becker, 1907, Pholeomyia ...... 108 atra (Duda, 1935), (Liodesma), Desmometopa ...... 99 atypica Sabrosky, 1983, Desmometopa ...... 97 Australimyza Hanison, 1953 ...... -> Australimyzidae 114 Irina Brake ENT. SCAND. SUPPL. 57 (2000) bakeri Aldrich, 1931, Milichiella ...... 105 beardsleyi Hardy & Delfinado, 1980, Leptometopa ...... 100 beckeri Kramer, 1920, Phyllomyza ...... 94 bimaculata Becker, 1907, Milichiella ...... 105 bisignata Melander, 1913, Milichiella ...... 105 blantoni Sabrosky, 1983, Desmometopa ...... 97 (Zetterstedt, 1848), (Milichia), ...... -> Odiniidae brevifacialis Deeming, 1976, Milichia ...... 103 brevirostris (de Meijere, 1909), (Prosaetomilichia), Milichia ...... 103 britannica Hennig, 1937, Madiza ...... 101 broersei de Meijere, 1946, Leptometopa ...... 100 cacahuamilpensis Henera, 1892, Pholeomyia ...... 108 Cacoxenus Loew, 1858 ...... -> Drosophilidae Carnus Nitzsch, 1818 ...... -> Carnidae cavernae de Meijere, 1914, Phyllomyza ...... 94 ciliata Hendel, 1919, Desmometopa ...... 97 ciliata (misidentification) Malloch, 1924, Desmometopa ...... 97 cinerea (Coquillett, 1900), (Ophthalmomyia), Milichiella ...... 105 cingulata Becker, 1907, Milichiella ...... 105 circularis Aldrich, 1931, Milichiella ...... 105 circularis Illingworth, 1929, Milichiella ...... 105 Cleptoneria Lioy, 1864, Madiza ...... 10 1 Cnemoplegas Enderlein, 1927, Leptometopa ...... 99 comans Sabrosky, 1959, Pholeomyia ...... 107 concavum (Becker, 1907), (Eccoptomma), Milichiella ...... 105 confusa (Cunan, 1934), (Desmomyza), Madiza ...... 101 cooperi Sabrosky, 1955, Eusiphona ...... 102 coquilletti (Hendel, 1907), (Hypaspistomyia), Leptometopa ...... 100 cornesi Deeming, 1976, Milichia ...... 103 Costalima Sabrosky, 1953 ...... 93 dampfi Sabrosky, 1959, Pholeomyia ...... 107 decora (Loew, 1870), (Lobioptera), Milichia ...... 103 decorior Steyskal, 1943, Pholeomyia ...... 107 dectes Collin, 1921, Milichia ...... 103 deserticola Ozerov, 1992, Xenophyllomyza ...... 96 Desmometopa Loew, 1866 ...... 96 Desmometopina Cunan, 1930, Leptometopa ...... 100 Desmomyza Cunan, 1934, Madiza ...... 101 Dicraeoptera Duda, 1932, Leptometopa ...... 100 dilatata Malloch, 1914, Phyllomyza ...... 94 dimidiata (Wiedemann, 1830), (Chlorops), Milichiella ...... -...... 105 dimorphic a Deeming, 1981, Enigmilichia ...... 102 discipalpis Papp, 1993, Desmometopa ...... 97 dispar (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 107 distinctipennis Deeming, 1981, Milichia ...... 103 donisthorpei Schmitz, 1923, Phyllomyza ...... 94 Eccoptomma Becker, 1907, Milichiella ...... 104 ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 115 e1ephas (Hendel, 1913), (A1drichiella), A1drichiomyza ...... 93 Enigmi1ichia Deeming, 1981 ...... 102 epitacta Hendel, 1914, Phyllomyza ...... 94 equitans (Hendel, 1919), (Neophyllomyza), Phyllomyza ...... 95 Eusiphona Coquillett, 1897 ...... 102 evanescens Sabrosky, 1983, Desmometopa ...... 97 excelsior (Becker, 1907), (Rhynchomilichia), Pho1eomyia ...... 107 eximia Papp, 1993, Madiza ...... 101 expansa Aldrich, 1925, Pho1eomyia ...... 107 fagicola Hendel, 1924, Phyllomyza ...... 95 farquharsoni Collin, 1921, Mi1ichia ...... 103 fasciventris (Becker, 1907), (Rhynchomilichia), Pho1eomyia ...... 107 flava Sabrosky, 1953, Eusiphona ...... 102 flava Sabrosky, 1958, Stomosis ...... 96 flaviceps (Enderle in, 1934), (Rhynchomadiza), Leptometopa ...... 100 flavicoxa Hendel, 1932, Desmometopa ...... 97 flavipa1pis de Meijere, 1914, Phyllomyza ...... 95 flavipa1pis Hendel, 1932, Milichiella ...... 105 flavipa1pis Sabrosky, 1983, Desmometopa ...... 97 jlavipalpis Seguy, 1932, Leptometopa ...... 101 jlavipes (Meigen, 1830), (Opomyza), Phyllomyza ...... 96 flavitarsis (Meigen, 1830), (Agromyza), Phyllomyza ...... 95 jlavitarsis (Meigen, 1830), (Opomyza), Phyllomyza ...... 96 jlavitarsis (Meigen, 1830), (Piophi1a), Madiza ...... 102 flaviventris Iwasa, 1997, A1drichiomyza ...... 93 flavoscutellata Malloch, 1925, Stomosis ...... 96 floridensis Sabrosky, 1983, Desmometopa ...... 97 fonnicae Schmitz, 1923, Phyllomyza ...... 95 fonnicophi1a Deeming, 1976, Mi1ichia ...... 103 formosus (Loew, 1844), (Milichia) Neoalticomems ...... -> Odiniidae ji-eyi (Hendel) in Frey 1936, (Eccoptomma), Milichiella ...... 105 frontale (Becker, 1907), (Eccoptomma), Milichiella ...... 105 fumicostata Deeming, 1981, Milichia ...... 103 fumipennis Malloch, 1934, Paramyia ...... 94 fuscogrisea (Seguy, 1933), (Neophyllomyza), Phyllomyza ...... 95 gigantea Deeming, 1979, Milichia ...... 103 glabra Fallen, 1820, Madiza ...... 101 glabrifrons Sabrosky, 1965, Litometopa ...... 101 glaucanota Sabrosky, 1983, Desmometopa ...... 97 gressitti Sabrosky, 1983, Desmometopa ...... 97 halteralis (Coquillett, 1900), (Desmometopa), Leptometopa ...... 100 Hemeromyia (Hendel, 1907) ...... -> Camidae Hendelimyza Duda, 1935, Phyllomyza ...... 94 hendeli Brake, n.n., Milichiella ...... 105 hirtipalpis Malloch, 1913, Phyllomyza ...... 95 Horaismoptera Hendel, 1907 ...... -> Tethinidae hungarica Papp, 1993, Paramyia ...... 94 116 Irina Brake ENT. SCAND. SUPPL. 57 (2000) hurdi Sabrosky, 1959, Pholeomyia ...... 107 Hypaspistomyia Hendel, 1907, Leptometopa ...... 99 ilaroensis Deeming, 1976, Milichia ...... 103 implicata (Becker, 1907), (Pseudomilichia), Pholeomyia ...... 107 inaurata Lamb, 1914, Desmometopa ...... 97 inconspicua de Meijere, 1916, Paramyia ...... 94 indecora (Loew, 1869), (Lobioptera), Pholeomyia ...... 107 indistincta Sabrosky, 1983, Desmometopa ...... 97 inimica Seguy, 1938, Neophyllomyza ...... 93 innominata (Williston, 1896), (Agromyza), Stomosis ...... 96 insecta (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 107 integra Becker, 1922, Milichia ...... 103 interfrontalis Sabrosky, 1965, Desmometopa ...... 97 javana de Meijere, 1911, Milichiella ...... 105 kandyensis Sabrosky, 1983, Desmometopa ...... 97 kaszabi Papp, 1976, Leptometopa ...... 100 lacteipennis (Hendel, 1913), (Madiza), Leptometopa ...... 100 lacteipennis (Loew, 1866), (Lobioptera), Mi1ichiella ...... 105 lacteiventris Malloch, 1931, Milichiella ...... 106 lasiae (Duda, 1935), (Hendelimyza), Phyllomyza ...... 95 latifrons Sabrosky, 1959, Pholeomyia ...... 108 latigena Sabrosky, 1983, Desmometopa ...... 98 latigenis (Hendel, 1933), (Hypaspistomyia), Leptometopa ...... 101 latipes (Meigen, 1830), (Agromyza), Leptometopa ...... 100 leanderi (Hendel, 1924), (Phyllomyza, subgenus Neophyllomyza), Neophyllomyza ...... 93 Leptometopa Becker, 1903 ...... 99 leptometopoides Sabrosky, 1983, Desmometopa ...... 98 leucogastra (Loew, 1861), (Milichia), Pholeomyia ...... 108 Leucopis Meigen, 1830 ...... -> Chamaemyiidae leucozona Bilimek, 1867, Pholeomyia ...... 108 Liodesma Duda, 1935, Desmometopa ...... 97 Liodesmometopa Duda, 1935, Desmometopa ...... 97 Litometopa Sabrosky, 1965 ...... 101 Lobioptera Wahlberg, 1847, Milichia ...... 102 longifacies Hendel, 1933, Pholeomyia ...... 108 longipalpis (Schmitz, 1924), (Neophyllomyza), Phyllomyza ...... 95 longirostris Hendel, 1931, Aldrichiomyza ...... 93 longiseta (Becker, 1907), (Milichia), Pholeomyia ...... 108 longiseta Hardy & Delfinado, 1980, Milichiella ...... 106 lucens Hendel, 1924, Phyllomyza ...... 95 lucidifrons Sabrosky, 1983, Desmometopa ...... 98 lucidula Becker, 1907, Milichiella ...... 106 ludens (Wahlberg, 1847), (Lobioptera), Milichia ...... 103 lute a de Meijere, 1914, Phyllomyza ...... 95 luteipalpis Malloch, 1914, Phyllomyza ...... 95 luteola (Coquillett, 1902), (Desmometopa), Stomosis ...... 96 luteus Aldrich, 1926, Microsimus ...... 93 ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae 117

Macromilichia Hendel, 1932, Pho1eomyia ...... 107 Macrurus Lioy, 1864, Leptometopa ...... 99 maculata (Meigen, 1830), (Milichia), Odinia ...... -> Odiniidae Madiza Fallen, 1810 ...... 101 Madizinae ...... 96 magnicomis Sabrosky, 1983, Desmometopa ...... 98 magnipalpis (Williston, 1896), (Phyllomyza), Neophyllomyza ...... 93 magnipalpus (elTor) Melander, 1913, Neophyllomyza ...... 94 mallochi Sabrosky, 1989, Leptometopa ...... 100 Mallochiella Melander, 1913, Leptometopa ...... 99 marginata (Mik, 1864), (Lobioptera), Milichia ...... 104 matilei Sabrosky, 1979, Leptometopa ...... 100 m-atrum (Meigen, 1830), (Agromyza), Desmometopa ...... 99 mcc1urei Sabrosky, 1964, Leptometopa ...... 100 medioCl"is Sabrosky, 1958, Milichia ...... 104 me1a1euca (Loew, 1863), (Argyrites), Milichiella ...... 106 melanderi Sabrosky, 1983, Desmometopa ...... 98 melania (Hendel, 1919), (Neophyllomyza), Phyllomyza ...... 95 Meoneura Rondani, 1856 ...... -> Camidae meridionalis Sabrosky, 1983, Desmometopa ...... 98 microps Lamb, 1914, Desmometopa ...... 98 Microsimus Aldrich, 1926 ...... 93 Mi1ichia Meigen, 1830 ...... 102 Milichiella Giglio-Tos, 1895 ...... 104 Mi1ichiinae ...... 102 mi1nei Steyskal, 1942, Phyllomyza ...... 95 lninuta Walker, 1865, Milichia ...... 103 minutissima (Wulp in Kertesz, 1897), (Agromyza), Neophyllomyza ...... 94 mira Coquillett, 1897, Eusiphona ...... 102 mixta Becker, 1907, Mi1ichia ...... 103 m-nigmm (Zetterstedt, 1848), (Agromyza), Desmometopa ...... 98 mongolica Papp, 1976, Phyllomyza ...... 95 montanum (Becker, 1907), (Eccoptomma), Milichiella ...... 106 morosa (Meigen, 1830), (Agromyza), Dizygomyza ...... -> Agromyzidae myopa Melander, 1913, Pho1eomyia ...... 108 mynnecophila (de Meijere, 1909), (Prosaetomilichia), Milichia ...... 103 myrmicola Sabrosky, 1953, Costalima ...... 93 nearctica Sabrosky, 1983, Desmometopa ...... 98 Neophyllomyza Melander, 1913 ...... 93 nigeriae (Duda, 1935), (Milichia), Mi1ichiella ...... 106 nigeriae Sabrosky, 1983, Desmometopa ...... 98 nigra (Williston, 1896), (Platophrymia), Desmometopa ...... 99 nigra Williston, 1897, Paramyia ...... 94 nigrella Cole, 1912, Milichiella ...... 105 nigricosta (Hendel, 1932), (Macromilichia), Pholeomyia ...... 108 nigripalpis de Meijere, 1914, Phyllomyza ...... 95 nigripes Malloch, 1931, Milichiella ...... 106 118 Irina Brake ENT. SCAND. SUPPL. 57 (2000) nigrohalteralis Sabrosky, 1983, Desmometopa ...... 98 nilotiea Becker, 1903, Desmometopa ...... 98 ni1ssoni Sabrosky, 1987, Leptometopa ...... 101 nitens (Loew, 1869), (Phyllomyza), Paramyia ...... 94 nitens Melander, 1913, Neophyllomyza ...... 94 nitidaBecker, 1907,Milichia ...... 103 nitida Hendel, 1911, Milichiella ...... 105 nitida (Walker, 1836), (Gymnopa), Milichiella ...... 106 nitidula Sabrosky, 1959, Pholeomyia ...... 108 niveipennis (Strobl, 1900), (Siphonella), Leptometopa ...... 101 niveipennis fascifrons (Becker, 1907), (Desmometopa), Leptometopa ...... 10 1 nudigena Sabrosky, 1983, Desmometopa ...... 98 nudipalpis Malloch, 1914, Phyllomyza ...... 95 nudiventris Becker, 1907, Milichiella ...... 106 obscura Sabrosky, 1959, Pholeomyia ...... 108 obscurifrons Sabrosky, 1983, Desmometopa ...... 98 Ophthalmomyia Williston, 1896, Milichiella ...... 104 orientalis Malloch, 1913, Mi1ichia ...... 103 orWia (Curran, 1927), (Mallochiella), Leptometopa ...... 100 pachymera Becker, 1908, Madiza ...... 102 pacifica Papp, 1984, Leptometopa ...... 101 pallida de Meijere, 1940, Phyllomyza ...... 95 palpalis de Meijere, 1914, Desmometopa ...... 99 pa1paris (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 108 palposa Zetterstedt, 1848, Milichia ...... 103 parafacialis Sabrosky, 1983, Desmometopa ...... 98 Paramadiza Malloch, 1913, Leptometopa ...... 99 Paramiliehia Malloch, 1913, Pholeomyia ...... 107 Paramyia Williston, 1897 ...... 94 Pareeeoptomma Duda, 1935, Milichiella ...... 104 parva (Macquart, 1843), (Milichia), Mi1ichiella ...... 106 patrizii Hennig, 1952, Milichia ...... 103 pecki Papp, 1984, Leptometopa ...... 101 pectoralis Hendel, 1932, Pholeomyia ...... 108 philippinensis Sabrosky, 1983, Desmometopa ...... 98 Pholeomyia Bi1imek, 1867 ...... 107 Phyllomyza Fallen, 1810 ...... 94 Phyllomyzinae ...... ,...... 93 pieta (Loew, 1861), (Milichia), Odinia ...... -> Odiniidae piscivora Malloch, 1926, Milichia ...... 103 Platophrymia Williston, 1896, Desmometopa ...... 96 pleuralis Sabrosky, 1983, Desmometopa ...... 98 poeci1ogastra Becker, 1907, Ulia ...... 109 politifacies Sabrosky, 1959, Pholeomyia ...... 108 Polphorina Enderlein, 1921, Madiza ...... 101 populi Steyskal, 1949, Milichiella ...... 106 posticata Becker, 1907, Milichia ...... 103 ENT. SCAND. SUPPL. 57 (2000) Phylogeny of Milichiidae II') postorbitalis Sabrosky, 1983, Desmometopa ...... 9X praeocellaris Hendel, 1932, Pho1eomyia ...... lOX praesecta (Becker, 19(7), (Rhynchomilichia), Pholeomyia ...... lOX Prodesmometopa Hendel, 1914, Leptometopa ...... 99 proectes Collin, 1921, Milichia...... 103 prominens (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 108 propeciliata Sabrosky, 1983, Desmometopa ...... 98 prosaetes Collin, 1921, Milichia ...... 104 Prosaetomilichia de Meijere, 1909, Milichia ...... 102 pseudodecora (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 108 pseudodectes (Seguy, 1933), (Milichia), Milichiella ...... 106 pseudoludens Papp, 1977, Milichia ...... 104 Pseudomilichia Becker, 1907, Pholeomyia ...... 107 Pseudopomyza Strobl, 1893 ...... -> Pseudopomyzidae Pseudorhiclloessa Malloch, 1914 ...... ->Tethinidae pubescens Becker, 1907, Milichia ...... 104 pulchra (Zetterstedt, 1848), (Milichia), Neoalticomerus ...... -> Odiniidae quadricomis Melander, 1913, Neophyllomyza ...... 94 quadrifasciata Hendel, 1932, Pholeomyia ...... 108 quercus Duda, 1935, Neophyllomyza ...... 93 Rhicnoessa Loew, 1862 ...... -> Tethinidae Rhodesiella Adams, 1905 ...... -> Chloropidae Rhynchomadiza Enderlein, 1934, Leptometopa ...... 100 Rhynchomilichia Hendel, 1903, Pholeomyia ...... 107 Risa Becker, 1907 ...... -> Risidae robertsoni (Coquillett, 1902), (Milichia), Pholeomyia ...... 108 rubricomis Schmitz, 1923, Phyllomyza ...... 95 rufifrons Becker, 1903, Leptometopa ...... 101 rujitarsis (Meigen, 1938), (Gymnopa), Madiza ...... 102 rufula (Frey, 1918), (Siphonomyiella), Stomosis ...... 96 saguaro Sabrosky, 1983, Desmometopa ...... 98 savannaticola Deeming, 1981, Milichia ...... 104 schineri Hendel, 1932, Pholeomyia ...... 108 schnusei (Becker, 19(7), (Pseudomilichia), Pholeomyia ...... 108 securicomis Fallen, 1823, Phyllomyza ...... 95 seguyi Hennig, 1937, Milichia ...... 104 semiaurata Sabrosky, 1958, Desmometopa ...... 97 silesiaca (Duda, 1935), (Hendelimyza), Phyllomyza ...... 96 simplicipes (Becker, 1907), (Desmometopa), Leptometopa ...... 101 sinaiensis Steyskal, 1966, Milichia ...... 103 singaporensis Kertesz, 1899, Desmometopa ...... 99 singaporensis: authors, in pati, not Kertesz, Desmometopa ...... 99 Siphonomyiella Frey, 1918, Stomosis ...... 96 smithi Aldrich, 1931, Milichiella ...... 106 solitaria (Lamb, 1914), (Eccoptomma), Milichiella ...... 106 sordida (Fallen, 1820), (Madiza), Desmometopa ...... 99 sororcula (Becker, 1907), (Rhynchomilichia), Pholeomyia ...... 109 120 Irina Brake ENT. SCAND. SUPPL. 57 (2000) speciosa Meigen, 1830, Milichia ...... 104 speciosa canariensis Becker, 1907, Milichia ...... 104 spinthera Hendel, 1913, Milichiella ...... 106 srilankae Sabrosky, 1983, Desmometopa ...... 99 Stenoporomyia Hendel, 1903, Milichiella ...... 104 stilbopleura Sabrosky, 1983, Desmometopa ...... 99 StOlllosis Melander, 1913 ...... 96 sumptuosa de Meijere, 1911, Milichiella ...... 106 superba Lamb, 1914, Milichia ...... 104 sylvicola Deeming, 1981, Milichia ...... 104 tamaricis (Bigot, 1884), (Milichia), ?Odinia ...... -> ?Odiniidae tarsalis Loew, 1866, Desmometopa ...... 99 tarsalis: authors, not Loew, Desmometopa ...... 99 tarsata (Zetterstedt, 1848), (Milichia), Phyllomyza ...... 95 tenebrosa Brunetti, 1924, Phyllomyza ...... 96 terminal is Sabrosky, 1983, Desmometopa ...... 99 Tethina Haliday, 1838 ...... -> Tethinidae tetragona Hendel, 1924, Phyllomyza ...... 96 texensis Sabrosky, 1959, Pholeomyia ...... 109 tiefii (Mik, 1887), (Lobioptera), Milichiella ...... 105 tosi Becker, 1907, Milichiella ...... 106 tricincta Becker, 1907, Milichiella ...... 106 tristicula Hendel, 1914, Desmometopa ...... 99 tristis Lamb, 1914, Milichiella ...... 106 Ulia Becker, 1907 ...... 109 unicolor (de Meijere, 1906), (Milichia), Milichiella ...... 106 urbana Malloch, 1913, Milichiella ...... 106 varipalpis Malloch, 1927, Desmometopa ...... 99 velutina Becker, 1907, Milichiella ...... 106 Vichyia Villeneuve, 1920, Neophyllomyza ...... 93 vidua Becker, 1907, Milichiella ...... 106 vittata Malloch, 1925, Stomosis ...... 96 vittata Sabrosky, 1982, Eusiphona ...... 102 vockerothi Sabrosky, 1961, Pholeomyia ...... 109 woldai Sabrosky, 1983, Desmometopa ...... 99 wulpi (Hendel, 1907), (Desmometopa), Neophyllomyza ...... 94 Xenophyllomyza Ozerov, 1992 ...... 96 Entomologica scandinavica, Supplement (Valid to I January 2002) US$ 1. Reports from the Lund University Ceylon Expedition in 1962. Vol. 1. 1971. XXXVI + 292 pp ...... 33 2. Palm, T.: Die skandinavischen Elateriden-Larven (Coleoptera). 1972,63 pp ...... 9 3. Nyholm, T.: Die nordamerikanischen Arten der Gattung Cyphon Paykull (Coleoptera). 1972. 100 pp ..... 11 4. Reports from the Lund University Ceylon Expedition in 1962. Vol. II. 1975. IV+ 239 pp ...... 30 5. Lindroth, C. H., Andersson, H., Bodvardsson, H. & Richter, S. H.: Surtsey, Iceland. The development of a new fauna, 1963-1970. Terrestrial invertebrates. 1973.280 pp ...... 28 6. Birket-Smith, S. J. R.: On the abdominal morphology of Thysanura (Archaeognatha and Thysanura s.st1'.). 1974.67 pp ...... 11 7. Moller Andersen, N.: The Limllogonus and Neogaris of the Old World with character analysis and a reclassification of the GelTinae (: GelTidae). 1975. 96 pp...... 14 8. Andersson, H.: Taxonomic and phylogenetic studies on Chloropidae (Diptera) with species reference to Old World genera. 1977.200 pp ...... 24 9. Heie, O. E.: Revision of the aphid genus Nasonovia Mordvilko, including Kakimia Hotles & Frison, with keys and descriptions of the species of the world (Homoptera: Aphididae). 1979. 104 pp...... 17 10. Srether, O. A. (Ed.): Recent developments in chironomid studies (Diptera: ). 1979. 150 pp ...... 25 11. Cederholm, L. (Ed.): Reports from the Lund University Ceylon Expedition in 1962. Vol. 3. 1982. 274 pp ...... 37 12. Huggert, L.: Revision of the West Palaearctic species of the genus Idris Forster s. I. (Hymenoptera, Proctotrupoidea: Scelionidae). 1979.60 pp ...... 10 13. Griffiths, G. C. D.: Studies on boreal Agromyzidae. XIV. Chromatomyia miners on Monocotyledones (Diptera: Agromyzidae). 1980.61 pp ...... 11 14. Srether, O. A.: Glossary of chironomid mOlphology terminology (Diptera: Chironomidae). 1980.51 pp. 17 15. Cederholm, L. (Ed.): Advances in insect systematics and phylogeny. 1981. 415 pp ...... 55 16. Srether, O. A. Orthocladiinae (Diptera: Chironomidae) from the British West Indies, with descriptions of Antillocladius n.gen., Lipurometriocllemus n.gen., COlllpterosmittia n. gen. and Diplosmittia n.gen. 1981. 46 pp ...... 11 17. Ashe, P.: A catalogue of chironomid genera and subgenera of the world including synonyms (Diptera: Chironomidae). 1983. 68 pp...... 19 18. Enghoff, H.: The milliped genus Cylidroiulus on Madeira - an insular species swarm (Diplopoda, Julida: Julidae). 1982. 142 pp ...... 26 19. Wiederholm, T. (Ed.): Chironomidae of the Holarctic region. Keys and diagnoses. Vol. 1. Larvae. 1983. 457 pp. Hardbound...... 80 20. Srether, O. A. & Sublette, J. E.: A review of the genera Doithrix n. gen., Georthocladius Strenzke, Paracizaetocladius Wlilker and Pseudorthocladius Goetghebuer (Diptera: Chironomidae, Orthocladiinae). 1983. 100 pp ...... 20 21. Fjellberg, A.: Arctic Collembola I - Alaskan Collembola of the families Poduridae, Hypogastruridac, Odontellidae, Brachystomellidae and Neanuridae. 1985. 126 pp ...... 20 22. Roth, L. M.: A taxonomic revision of the genus Blattella Caudell (Dictyoptera, Blattaria: B1attellidac). 1985.221 pp...... '10 23. Loken, A.: Scandinavian species of the genus Psithyrus Lepeletier (Hymenoptera: Apidae). 1984. 45 pp. I () 24. Lomholdt, 0.: A reclassification of the lan'ine tribes with a revision of the Miscophini of southern Africa and Madagascar (Hymenoptera: Sphecidae). 1985. 183 pp ...... :1.'\ 25. Nielsen, E. S.: A taxonomic review of the adelid genus Nematopogon Zeller (: Incurvarioidea). 1985. 66 pp...... 12 26. Hansson, c.: Taxonomy and biology of the palearctic species of Clllysocharis Forster, 1856 (Hymenoptera: Eulophidae). 1985. 130 pp ...... )() 27. Gardenfors, U.: Taxonomic and biological revision of Palearctic Ephedrus (Hymenoptera: Braconidae, Aphidiinae). 1986.95 pp ...... 1'/ 28. Wiederholm T. (Ed.). Chironomidae of the Holarctic region. Keys and diagnoses. Vol. 2. Pupae. 1983.482 pp. Hardbound ...... HI) 29. Srether, O. A. (Ed.) A conspectus of contemporary studies in Chironomidae (Diptera). Contributiolls from the IXth Symposium on Chironomidae, Bergen, Norway. 1987.393 pp. Hardbound...... SH 30. Cederholm, L. (Ed.): Reports from the Lund University Ceylon Expedition in 1962. Vol. IV. 1988. 160 pp ...... 1:, 31. Hansson, C. Revision of the New World species of Cluysocharis Forster (Hymenoptera: Eulophidae). 1987.86 pp ...... :...... 15 32. Bengtson, S.-A. & Enckell, P. H. (Eds.): Island terrestrial invertebrates: Species composition, dispersal and faunal history. 1988.88 pp ...... 23 33. Ronquist, F. & Nordlander, G. Skeletal morphology of an archaic cynipoid, Ibalia rufipes (Hymenoptera: Ibaliidae). 1989.60 pp ...... 20 34. Wiederholm, T. (Ed.): Chironomidae of the Holarctic region. Keys and diagnoses. Vol. 3. Male imagines. (Diptera). 1989. Hardbound. 538 pp ...... 97 35. Srether, O. A.: A review of the genus Limnophyes Eaton from the Holarctic and Afrotropical regions (Diptera: Chironomidae, Orthocladiinae). 1990. 135 pp ...... 40 36. Scharff, N.: Spiders of the family Linyphiidae from the Uzungwa mountains, Tanzania (Araneae). 1990.95 pp ...... 40 37. Olafsson, E. Taxonomic revision of wester Palaearctic species of the genera Scatella R.-D. and Lamproscatella Hendel, and studies on their phylogenetic position within the subfamily Ephydrinae (Diptera, Ephydrinae). 1991. 100 pp ...... 45 38. Hansson, C. A catalogue of the Chalcidoidea (Hymenoptera) described by C. G. Thomson, with a checklist of Swedish species. 1991. 70 pp...... 25 39. Muona, 1. A revision of the Indomalesian tribe Galbitini new tribe (Coleoptera, Eucnemidae). 1991.66 pp ...... ;...... 25 40. Enghoff, H. Dolichoiulus - a mostly Macaronesian multitude of millipedes. With a description of a related new genus from Tenerife, Canary Islanmds (Diplopoda, Julida, Julidae). 1992. 158 pp ...... 55 41 Tjeder, B.t & Hansson, Ch. The Ascalaphidae of the Afrotropical Region (Neuroptera). 1992.240 pp ... 70 42. Baranowski, R. Revision of the genus Leiodes Latreille of North and Central America (Coleoptera: Leiodidae). 1993. 149 pp ...... 55 43. Nolte, U. Egg masses of Chironomidae (Diptera). A review, including new observations and a preliminary key. 1993.75 pp ...... 30 44. Muona, J. Review of the phylogeny, ciassification and biology of the family Eucnemidae (Coleoptera). 1993. 133 pp ...... 50 45. Pape, Th. The World Blaesoxipha Loew, 1861 (Diptera: Sarcophagidae). 1994.247 pp ...... 70 46. Ronquist, F. Phylogeny and classification of the Liopteridae, an archaic group of cynipoid wasps (Hymenoptera). 1995.74 pp ...... 30 47. Scheller, U. First record ofPauropoda (Myriapoda) from Sierra Leone. 1995.54 pp ...... 25 48. Srether, O. A. & Wang, X. Revision of the genus Paraphaenocladius Thienemann, 1924 of the world (Diptera: Chironomidae, Orthocladiinae). 1995.69 pp ...... 30 49. Hansson, Ch. Taxonomic revision of the Nearctic species of Omphale Haliday (Hymenoptera: Eulophidae). 1996.78 pp ...... 35 50. Soli, G. E. E. On the morphology and phylogeny of , with a revision of Coelosia Winnertz (Diptera, ). 1997. 137 pp ...... 42 51. Enghoff, H. (Ed.) Many-legged -- A collection of papers on Myriapoda and Onychophora. Proceeding of the 10th Int. Congress of Myriapodology, Copenhagen 29 July-2 August 1996. 1997. 329 pp ...... 80 52. Hedqvist, K-J. Bark enemies in Sweden. II. Braconidae (Hymenoptera). 1998. 87 pp ...... 38 53. Johanson, K A. Phylogenetic and biogeographic analysis of the family HeJicopsychidae (Insecta: Trichoptera). 1998. 172 pp ...... 60 54. Lundqvist, Lars. Taxonomic revision of the genus Dinogamasus Kramer (Acari: Mesostigmata: Laelapidae). 1999. 109 pp ...... , ...... 40 55. Komai, Furumi. A taxonomic review of the genus Grapholita and allied genera (Lepidoptera: Tortricidae) in the Palaearctic region. 1999.226 pp ...... 70 56. Adam, J. I. & Srether, O. A. Revision of the genus Nilothauma Kieffer, 1921 (Diptera: Chironomidae). 1999. 106 pp ...... 40 57. Brake, I. Phylogenetic systematics of the Milichiidae (Diptera, Schizophora). 2000. 120 pp ...... 45

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